Beruflich Dokumente
Kultur Dokumente
m
r* tr
o m a
ANATOMY
OF THE
CORD
ANATOMY
OF THE
CORD
BY
HARRIS
E.
SANTEE, M.
D.,
Ph.D.
PROFESSOR (.F ANATOMY IN THE COLLEGE OF PHYSICIANS AND SURGEONS, MEDICAL DEPARTMENT, UNIVERSITY OF ILLINOIS; PROFESSOR OF ANATOMY IN JENNBR MEDICAL COLLEGE, CHICAGO; MEMBER OF ASSOCIATION OP AMERICAN ANATOMISTS.
WITH
128 ILLUSTRATIONS, 33
IN
COLORS
PHILADELPHIA
P.
WALNUT STREET
1907
&
Co.
PREFACE
THE
receive
may
from
his
colleagues
the
anatomy
human
brain and
To do
was
have been gleaned from many possible, from the original sources.
is
presented
description from the gross structures to the constituent neurones in proceeds each successive region. Wherever the embryology will assist the
The
student to comprehend the adult forms, the development is briefly given in the text but a special chapter is also devoted to embry;
which presents a concise and connected statement of the development of the entire brain and spinal cord.
ology,
The
their afferent,
Particular emphasis is laid the origin, course, termination and function of conduction upon paths as they are met in the regular study, and the more important and better known of these paths are summed up in a
final
and
efferent connections.
Function
is
everywhere
given in connec-
The
BNA
tion, the
employed.
lectures, every student is expected to brain in the laboratory, exposing, studying and sketching every macroscopic structure as it occurs in the work;
dissect the
human
VI
PREFACE.
and, then with the microscope, examine the minute structure and For these purposes the picture the histology of the same parts.
class should
be taken in small
brain.
to four students,
sections, divided into groups of two and each group should be provided with a well
It is desirable that
hardened
human
receive a well stained microscopic section of every important part. This is, however, often impossible; and the instructor may get
along with considerable satisfaction, by having the students exchange, if he has but a few sets of slides.
The author
literature
and
"Development of The Human Body," Barker's "BNA" and " Nervous System," the work of Dr. Alfred W. Campbell, Histological Studies on the Localization of Cerebral Func" the "Text-book of Anatomy" and "Memoirs of D. J. tion," R. S., and the fourth edition of Morris's " Human Cunningham, F.
Barker's "
especially useful.
my
colleague, Prof.
my Wm.
appreciation of the kindly assistance of T. Eckley, M. D.; and to say that the
new illustrations in this book belongs to my friend and pupil, Mr. Zan D. Klopper, of Chicago, who sketched the original drawings from my own specimens. My thanks are
artistic
merit of the
also
due
favors
and
courtesies
select illustrations
little monograph, for many shown me, particularly for allowing me to from Gordinier, McMurrich and Morris, works
published by them.
HARRIS E. SANTEE.
TABLE OF CONTENTS.
CHAPTER
I.
Dura Mater
Structure
of the Brain
i i
i
and
relations
Processes
Sinuses
2-5
5
6-7
7
7
Nerves
Contrasted with dura of spinal cord Arachnoid of the Brain
:
8
8
Structure
Relations, subarachnoid spaces. Vessels and nerves
8-9
9
9
and
relations
9-10
10
and veins
10-13
13
Nerves
Contrasted with pia of spinal cord Blood Supply of the Brain:
Carotid and vertebral arteries
13
14-30
14
14-2 1
and branches
14-15
15-18
16
16-17
17
and anterior
17-18 18-21
Vlll
TABLE OF CONTENTS.
PAGE
Antero-median ganglionic
arteries
18 18
18-21
21
2
The Veins
of the
Cerebrum:
1-24
21
1-22
22-23
22
Medial
Inferior
22-23
23
Lymphatics of cerebrum
B. Circulation of the Rhombencephalon
:
23-24
24-30
24
The medulla oblongata B2. The pons (Varolii) 63. The cerebellum
Bi.
Superior cerebellar artery Anterior inferior cerebellar artery Posterior inferior cerebellar artery
Internal cerebellar veins
24-27
27-30
2
7-28
29 29
29
29-30
29
29-30
30
Lymphatics of cerebellum
Table
Embryologic Divisions of the Brain Components of cerebrum
I.
3~33
33 33
Components
of
rhombencephalon
CHAPTER
II.
34 34
the ventricles
34~35
35~36
36-41
Anterior area
36-37
Middle area
Posterior area
37~38
38-41
TABLE OF CONTENTS.
Roots of the Twelve Cerebral Nerves:
Nuclei, genetic and terminal Olfactory nerves
IX
PAGE
42-49
42
42
Optic nerve
42
Oculo-motor nerve
Trochlear nerve
42-45
45 45 45
45-46
45-46
46
Intermediate nerve
Acustic nerve
46
46-49
49
49
CHAPTER
III.
THE CEREBRUM.
Subdivisions:
Enid -brain
Inter-brain
50 50 50
Mid-brain
SECTION
I.
THE
FORE-BRAIN OR.PROSENCEPHALON
50-144
5 I- 98
and sulcus
52 52
52
5 2 ~73
5 2 ~55
55
55-56
56
56-59
59~73
59-63
Frontal lobe,
and sulci Parietal lobe, its gyri and sulci Occipital lobe, its gyri and sulci
its
gyri
63-67
67-69
TABLE OF CONTENTS.
PAGE
Temporal
and
sulci
69-7 1
70 70-71 7 I- 73
The
73-85
73-74
74~77
Rhinencephalon
Olfactory lobe
Olfactory bulb
Olfactory tract and
78 79
79
79
Chorioidal
fissure
fissure
79-82 80
Hippocampal
80
80-81
81 81
81
Ectorhinal sulcus
Fissura collateralis
Inferior temporal sulcus
Gyrus Gyrus lingualis Limbic lobe, inferior part Gyrus hippocampi and uncus
Dentate fascia
r
fusiformis
81
81-82 81-82
82
Hypothalamus
Pars optica hypothalami
'.
82-85
82-84
83
Lamina
cinerea terminalis
83-84
84
84
82
and 84
84 84-98 86 86 86-89
(calloso-marginal)
sulcus
89
Calcarine fissure
89-90
90
Hippocampal
fissure
TABLE OF CONTENTS.
Chorioidal fissure
Collateral fissure
XI PAGE
90
Ectorhinal sulcus
Inferior temporal sulcus Gyri of medial and tentorial surface
9~93 9~93
93
93-98
93~97
Gyrus fornicatus
Gyrus cinguli Gyrus hippocampi Uncus Lobus pyraformis Limbic lobe, rhinencephalon
..
93~94
94~97
94-97
97
97
97
marginalis)
97
97 97
97
Cuneus
Gyrus lingualis Gyrus fusiformis
97 97
Summary
of lobes of the
cerebrum
98 98
Neopallium
Rhinencephalon
(archipallium)
,
98 98
Corpus striatum
Interior surface of the fore-brain
Motor
fibers fibers
100-104
occipital parts
103
Motor
fibers
103
Common
103-104
104
104-108
107 107 107
Upper
surface
107-108 108
108-109
109
Body
of fornix
Xll
TABLE OF CONTENTS.
PAGE
Crus
fornicis
fornicis
Columnae
no no
111
Septum pellucidum
Fifth ventricle
111-112
its
boundaries.
Lentiform nucleus
Nucleus caudatus
Stria terminalis
16-119
119
119 120
Thalamus
Chorioid plexus of lateral ventricle Anterior horn of the lateral ventricle
Posterior horn
Inferior
1
20-1 23
123
horn
collaterale
its
123-127
124
Trigonum
Chorioid
Hippocampus,
digitations
124
124
epithelium.
127-144
132
132
132-135
135-136 136-139
139
-.
139-143
".
Surfaces
140
and
inferior..
140-143
143 143
143
143-144
:
SECTION
II.
144-164
Surfaces
144-146
147-151
148 148
Temporo-pontal
Pyramidal tract
tract
148-151
151
Fronto-pontal tract
TABLE OF CONTENTS.
Substantia nigra.
Xlll
PAGE
151
Tegmenta
Cerebral aqueduct (Sylvii)
152-163
152
152-155
155
155-163
155
Tegmental decussations
Tracts of fibers in the tegmentum
155-163
Medial
bundle
155-158
158-159
159-161
Medial
Lateral
fillet fillet
superior
fillet.
159-160 160-161
161-162
162
162 162
Spino-thalamic tract.
Brachium conjunctivum
Rubro-spinal tract
Olivary fasciculus
163
163-164
163
Colliculus inferior
163-164
-
164 164
164-241
167-1 73
167 167 168
The neurone
Dendrites
Axone
Myelin sheath
168 168
Types
of neurones (three)
(ist, 2d, etc.)
Orders of neurones
168-171
171
Functions of neurones
171-172
172-173
173
Epiblastic
173-174
173-174
174
174-204
XIV
TABLE OF CONTENTS.
PAGE Cortical or cerebral localization
177-183
177
Motor
area, emissive
178
178
178 181
181 181
Optic center
Olfactory and gustatory centers
Naming
center
181
18 1
182
182-183
183
and
fiber lamination
183-204
184-187 184 and 187
184 and 187 184 and 187-188 184 and 188 184 and 188-191 184 and 191
192
.
Layer of
stellate cells
192
192-204
192-195
Olfactory cortex
195-204
195-196
196-199
199
Olfactory bulb
Uncus hippocampi
Nucleus amygdalae Subiculum
Fascia dentata
olfactorium, etc
199
200
200-203
203
Trigonum
204
204-221
Corpus striatum
Centrifugal fibers
Centripetal fibers
204-208
207-208 208
208-217 208-211
Thalamus
Thalamic nuclei..
.
TABLE OF CONTENTS.
White matter
Cortical
of the thalamus
XV
PAGE
211-217
fillet
216-217
217 217 217 218
Red nucleus...
Nucleus hypothalamicus Metathalamus
Lateral geniculate body
218
218
218-219
Nucleus
Hypothalamus
Pars optica Pars mammillaris
221-224 221-222
221
221-222
222
222-224
223
223
224
224-234
225-231
225 225
motor
fibers
Intermediate tract
Fronto-pontal tract
Tempora-pontal
Pyramidal
tract
tract
225-226
226-231
227 228
fibers
228
'
228-231
231
231-234
231
Medial
fillet,
functivum
Olfactory projection fibers
Cortical
fillet
232
(common
sensory)
232-233
233
XVI
TABLE OF CONTENTS.
PAGE
Auditory
Lateral
fillet
233
and brachium
inferius
233
233
Temporo-thalamic radiation
Occipito-thalamic radiation (optic)
234
Commissural
fibers of
cerebrum
234-236
234
Corpus callosum
Anterior commissure
234-235
236
Commissura hippocampi
Association fibers of cerebrum
236-241
236-238
238-241
238
Long
association fibers
of gyrus fornicatus
Cingulum
Fornix
238-240 240
240
240-241
241 241
Uncinate fasciculus
Superior longitudinal fasciculus
Inferior longitudinal fasciculus
Fasciculus occipito-frontalis
Perpendicular fasciculus
CHAPTER
IV.
THE RHOMBENCEPHALON.
SECTION
Function
Divisions
:
I.
THE CEREBELLUM:
242-266
342 242
242-243
243
244
Medullary Body:
Inferior medullary velum Brachia conjunctiva
244-247
244
245
246
246-247
247
247-248
248-25 1
248-249
248 249
TABLE OF CONTENTS.
Predeclivil sulcus
XV11 PAGE
249 249
Postdeclivil sulcus
Lobes of superior surface Lobus lingulae Lobus centralis Lobus culminis monticuli Lobus declivis monticuli Lobus folii vermis
Inferior Surface of Cerebellum:
Sulci of lower surface
250-251
250
250
250
251 251
251-256
252-253
252
Postnodular sulcus
Prepyramidal sulcus
Postpyramidal sulcus
Midgracile and postgracile Lobes of lower .surface
252-253
253
253
253-256
253-254
254
254-255
255-256
Gray Matter
of the Cerebellum
256-260 256-258
256
257.
Superficial layer
and
stellate cells
257
257-258
Deep
258-260
258-259 260
cells
Function of
stellate,
260
Neuroglia of cerebellum
260
260-262
260
26.0-261
Nucleus dentatus
261
262-266 263-265
263
263-264
264
264-265
XV111
TABLE OF CONTENTS.
PAGE
Commissural
fibers
265
Association fibers
265-266
(VAROLII)
SECTION
II.
THE PONS
Surfaces:
269-270
271-272
271 271
Transverse
Deep
271-272
272
Trapezoid body
Longitudinal fibers of the pons Ventral
272-278
272
Dorsal
272-278
fillet
fillet
Medial
Lateral
and superior
fillet
273-274
272 272
tract
Spino-thalamic tract
272-276
276
276
276
276-277
277 277
277
Gray Matter
277-284
277
278-284
278-279
279
279
279-280 280
281 281 281
Genetic (motor)
Terminal (sensory)
Nucleus of abducent nerve
281-282
282
Nucleus of
facial nerve..
TABLE OF CONTENTS.
Salivary nucleus Vestibular nucleus of auditory
XIX
PAGE
283
283
Lesions in pons
283
. .
SECTION
III.
2-84-313
Origin
Ventricle
-. .
285 285
Surfaces
286-290 286
286
Anterior
surface
286-287
287-288
287
287-288
Restiform body
288-290 288
of fourth ventricle
Roof epithelium
289
290
White matter
of medulla
290-302
291
Substantia reticularis
Raphe
Transverse fibers
291
291-292
291
291-292
292 292
decussation
Cerebello-olivary fibers
Dorso-ventral fibers
Anterior external arcuate
292-293
'292
293
Roots of eighth
293-298 294
295
296
297
298-300
298
tract
298
299
tract
Rubro-pinal
tract
300
300-302
XX
TABLE OF CONTENTS.
PAGE
300
300
301 301
Tractus
solitarius
Gray matter
of the medulla
fibers
302-3 13
302
nucleare)
303-310
303
303-304
Nucleus ambiguus
Nuclei
alae cincereae
Nucleus tractus
Vestibular nuclei
solitarii
307-308
309-3 10
3 10-3 13
gracilis
310-311 310-311
312
3 I 3~3 2 5
SECTION IV.
Boundaries
Floor of fourth ventricle
Colliculus facialis
313-314
314-318
317 317
317 317 317
Area acustica
Origin of cerebral nerves
320
320-325
320-321
Sensory Nerves and Roots Table III. Motor Nerves and Roots
Table
II.
321-322
Terminal nuclei
322-324
322 322
.
Common
Cortical
sensory
and
reflex connections
322-324
324
TABLE OF CONTENTS.
Genetic nuclei
.
XXI
PAGE
324-325
reflex connections
Cortical
and
324-325
326
326-329
3 2 9~33
:
Blood Supply of the Spinal Cord Spinal arteries, anterior and posterior
Fissural or centrifugal
33~33 2 33~33 2
331
33 I- 33 2
Centripetal
332
332 332 332
Root and
fissural veins
Lymphatics
332
333
333~335
333~334
334~335
337
Lumbar enlargement
Sixth ventricle (canalis centralis spinalis) Fissures of the cord:
337~33&
337
median
fissure
337
337~33&
338 338
Anterior root-line
(s.
lateralis anterior)
338-351
339
339
339
:
Gray
crescent
Anterior columna
Cells of anterior
columna
XX11
TABLE OF CONTENTS.
PAGE
340
340 340
340, 343-344
Medial column
Lateral column
Cortical connection
343
Reflex mechanism
343~344
344
lateralis
344-347
(efferent
column
of
cell-bodies
sympathetic neurones)
Posterior
344-347
columna Neurones of head of posterior columna Nucleus dorsalis (Clarki) terminal nucleus
pathetic)
347-349 347-348
of
sym348-349
350 350
350
350
351-362
351 351
35 1 ~35 2
352
. .
352
352 352
352 352
Funiculus
lateralis
Funiculus posterior
Ascending
tracts tracts
Descending
Mixed
tracts
352
35 2 ~353
Embryological method of locating tracts Pathological and experimental method Tracts of the Spinal Cord
Antero-lateral fasciculus proprius
353
354-362
354 354
355
355
tract
356
3 56-3 5 7
tract
356-357
357
TABLE OF CONTENTS.
(Posterior) Cerebello-spinal tract
XX111
PAGE
357~35&
357~358
358
358
359 359
359~36o
360-361
361 361
Descending
tracts
Descending postero-medial tract (comma, oval, septomarginal and median triangular tract) Descending postero-lateral
Posterior fasciculus proprius
tract
Cornu-commissural tract
Lesions of posterior columns
362
362
362-365
363 363
363 363
363
360-364
364-365
364
364 364
364-365
365
365
365
CHAPTER
I.
VII.
TRACING OF IMPULSES.
Efferent, or
Motor Paths:
366-374
366-369
366 369
Through
cerebral nerves
Cerebro-pontal paths
Fronto-pontal
369-370
370
XXIV
TABLE OF CONTENTS.
PAGE
Temporo-pontal
Intermediate
Spinal and cerebral Paths through red nucleus
370
370 370
37~373 37~373
373 373
brachium conjunctivum
tract
Medial ponto-spinal
373
373
II.
Lateral ponto-spinal tract Short fiber paths in formatio-reticularis Afferent, or Sensory Paths, General Sensations:
Tactile, muscular, pain,
(I).
373~374
377-383
377
skin,
377-38i
Through
377~378
378 378
Direct route
Indirect route
Through
(II).
cerebral nerves
tactile
Muscular and
378-381
..
381 381
Through
(III).
impulses
tract
tract
..
381-382
Through spino-thalamic tract Through ascending anterior cerebello-spinal Through cerebral nerves and spino-thalamic
Short fiber paths Afferent Paths Special Sensations:
Olfactory path
381-382 381-382
382
382-383
383-390
383-384
384-3 8 5
Optic path
385-389
386
386-389
389 389
III.
389-390
390^394
390
Reflex arcs
(1) Spinal reflexes
39-39 2
390
391-392
39 2 ~393
Cerebral reflexes
TABLE OF CONTENTS.
Spino-cerebral reflexes
XXV
PAE
(3) (4)
393
Cerebro-spinal reflexes
393~394
393
Respiratory reflexes
Equilibrium reflexes
Pupillary reflexes
393
394
395
Ependymal
Indifferent cells
397~398
398
Neuroblasts, neurones
Spongioblasts, neuroglia
398-399
399-400
400-401
: .
.'
Neural Crest
400 400
400-401
401
THE
BRAIN:
Flexures
401-426
402 403
Table IV. Brain Vesicles and Derivatives Table V. Secondary Brain Vesicles
Telencephalon (end-brain) Table VI. Derivatives of
Optic
vesicle,
403-404
404-415
404-405
405
405-415 406-409
409-410 410-411
411
41 1-412
Fornix
Stria terminalis
412
412
Internal capsule
412-413
Anterior commissure
413-414
Corpus callosum
414
XXVI
TABLE OF CONTENTS.
PAGE
Septum pellucidum
Pars optica hypothalami
415
415
415-417
415 416
417 417
Dorsal lamina
Ventral lamina and floor-plate
417-418
417-418 418-424
419
418-421
419-420 420-42 1
42 1
421
42 1-422
422-426
423
External surface
423-424 424
424-425
425
Ependymal
layer.
Mantle layer
Neuroglia layer, marginal velum
425-426
THE
SPINAL CORD:
Ventriculus terminalis
426-430 426
426 426 426
Filum terminale
Cauda equina
Meninges
Zones, ventral and dorsal
Histologic layers
426-427
427-428 428
428-429
429
Ependymal
layer
429-430 429-430
430
fissure
median
430 430
430
LIST
FIG.
OF ILLUSTRATIONS.
PAGB
1.
showing falx
a part of the tentorium cerebelli and the sinuses of the dura mater. (After Morris's Anatomy)
2.
Upper
and
cer. .
(After Morris's
Anatomy)
3.
Sinuses of the dura mater in the base of cranium, etc. Morris's Anatomy)
(After
5
4.
Coronal section of meninges showing falx cerebri, superior sagittal sinus and the arachnoid granulations. (Gordinier after
Key and
Retzius)
5.
6.
Middle meningeal artery inside the cranium. (After Morris's Anatomy) Diagram of pia mater and arachnoid, showing subarachnoid
spaces.
(After Morris's
Anatomy)
Fornix turned back to
7.
8.
show the chorioid tela of third ventricle. (Original), Roof and lateral walls of fourth yentricle and its chorioid
plexus.
(After Morris's
n
13
Anatomy)
branches.
and
its
The
base of the
15
(After
brain.
10. Arterial circle
(After Morris's
(Willisi)
n. Middle
12
19
(After Gordinier
from Quairi)
Anterior and posterior cerebral arteries.
13. Arteries of the 14.
22 25
27
(After Spalteholz).. (Gordinier after Duret) ... section of embryonic brain of third month. (After
medulla oblongata.
28
(After Morris's Anat-
31
vesicles.
16.
(After Morris's
Anatomy)..
35
17.
Diagrammatic horizontal section of vertebrate brain, showing vesicles and ventricles. (After Morris from Huxley)
xxvii
36
XXV111
FIG.
LIST OF ILLUSTRATIONS.
PAGE
18.
Diagrammatic median section of vertebrate brain showing vesicles, ventricles and olfactory diverticulum. (After Morris from Huxley)
(Original)
37 39
43
47
20.
21
.
(Original)
22. Latero-superior
53
23.
The convex
sulci.
and
57 61
24.
(Original)
island (Reili) and the superior surface of the temporal lobe, arachnoid granulations,
away
to
show the
etc.
(Original)
65
26.
Right
75
cut away.
(Original) (Original)
27.
28.
The median
sulci.
87
Medial surface of
and
91
(Original)
Gyri on medial surface of hemisphere. (Original) Transverse section of the brain, directed from the pons ob30.
29.
95
upward and forward, showing internal capsule, corpus callosum, ganglia and ventricles of the foreliquely
brain.
(Original)
101
and the
showing
33.
island.
(Original)
105
pus, inferior
horn of
(Original).
.-..
113
Diagram
(Original)
115
hemisphere cut
117
away
to
expose
it.
(Original)
corpus callosum, showing lateral ventricles, septum pellucidum, fornix and transverse temporal
inal)
36. Horizontal section of
gyri.
121
cerebrum
LIST OF ILLUSTRATIONS
FIG.
XXIX
PAGE
and
lateral ventricles.
(Or125
and
internal cerebral
(Original)
section of left cerebral hemisphere cutting the
129
39.
The
splenium and showing the posterior horn and the floor horn of the lateral ventricle. (Original) 131 inferior' and posterior horns of the lateral ventricle, shown
of the inferior
by removal
(Original)
133
40. Horizontal section of cerebrum through genu and below splenium of corpus callosum, fornix and chorioid tela turned
(Or-
137
of the ventricles.
Diagrammatic.
141
(Original)
42. Transverse
43.
The
(After Told. Morris's Anatomy) 145 region of the mid-brain showing pulvinar of the thalamus, the geniculate bodies, the corpora quadrigemina and
(Original)
The
dorsal or posterior aspect of the inter-brain, the the pons and the medulla. (Original)
mid -brain,
149
(Orig-
inal)
153
157
(Original) ..
48. Section of the mid-brain cutting the inferior colliculi of the cor-
pora quadrigemina.
49. Varieties of neurones in the
(Original)
human
nervous system.
50.
Motor neurone.
51.
An
Morris's Anatomy) 165 169 (After Barker's Nervous System) efferent neurone and an afferent neurone. (After Brubaker's Physiology)
171
in the spinal cord.
52.
172
XXX
FIG.
LIST OF ILLUSTRATIONS.
PAGE
53.
Neuroglia
54.
and ependyma cells of the spinal cord. Lenhossek, Gordinier's Nervous System) Cortical areas on convex surface of cerebral hemisphere.
cells
(After
173
(Or-
iginal)
55. Cortical areas
175
bral hemisphere.
56. Cortical areas after C.
(Original)
179
of cerebral
Convex surface
Medial and
hemisphere.
(Brubaker's Physiology)
182
tentorial surface
K. Mills.
of cerebral hemisphere.
58. Cell
(Bmbaker's Physiology)
area.
183
and
central
bell)
gyms.
The motor
(After
A W. Camp.
185
59. Cell
and
central
common
sensory area.
(After
189 Receptive
60. Cell
and and
visual area.
61. Cell
formis).
A. W. Campbell) 193 uncus hippocampi (lobus pyraThe area of smell. (After A. W- Campbell) 197
hippocampal region. (After Edinger) 201
(Gordinier after
Van
203
Gehuchteri)
64. Horizontal section of the
cerebrum through genu and below of corpus callosum, fornix and chorioid tela splenium turned back to show inter-brain and third ventricle.
(Original)
205
show geniculate
66.
nucleus amygdalae, etc. (After Morris's Anatomy) 207 Transverse section of the brain in the line of the pyramidal
tracts,
showing basal ganglia, internal capsules, corpus Viewed callosum, lateral and third ventricles, etc.
from
67.
front.
(Morris's
(Original)
Anatomy
after Toldt)
209
213
The
optic path.
68. Section of
colliculi
(Original)
69. Section of the mid-brain cutting the inferior colliculi of the cor-
pora quadrigemina.
70. Horizontal
(Original)
and
much
226
enlarged.
(Original)
LIST OF ILLUSTRATIONS.
FIG.
XXXI
PAGE
71. 72.
Diagram
(Original}
227
diagram showing motor and sensory paths, motor red, (After Gordinier's Central Nervous sensory blue.
73.
System) 229 Transverse section of cerebrum, cutting corpus callosum, anterior commissure and optic chiasma. Viewed from
front.
Commissural
fibers.
(Morris's
Anatomy
after
Toldf)
74.
-.235
Diagram
(Gordinier and Quain after Meynert) 237 Fasciculus occipito-frontalis. Stria terminalis and fasiculus 75. uncinatus. (Gordinier after Dejerine) 239 Su76. Dorsal view of inter-brain, mid-brain and cerebellum.
perior surface of cerebellum.
77. Anterior aspect of cerebellum.
78. Dissection of
(Original)
243
(Original)
245
79.
rhombencephalon to show brachium conjunctivum, brachium pontis and corpus restiforme. (Gordinier, Sappey after Hirschjeld and Leveille) 246 Median section of cerebellum, pons and medulla. (Original) 247
(Original)
252
(Or-
255
made
Diagrammatic.
border.
ten)
(Cunningham
258
Diagrammatic.
(Gordinier after
Van Gehuch259
and brachia
261
(After
(Morris's Anatomy after Toldf) conjunctiva. Anterior aspect of mid-brain, pons and medulla. 85. Morris's Anatomy)
86.
267
(Morris's
Anatomy
270
275
(Original)
bellum.
89.
(Original)
Diagram
Anatomy)
90. Section of embyronic medulla,
280
embryo measuring
Minot
after
9.1
mm.
in
length.
(Gordinier and
His)
285
XXX11
FIG.
LIST OF ILLUSTRATIONS.
.
PAGB
its
91.
Roof and
and
chorioid
(After Morris's Anatomy) 289 medulla oblongata near the pons. (Original) 295 (Or93. Section of the medulla oblongata at the middle of olive.
plexuses.
92. Section of
iginal)
94. Section of the
297
fillet
decussation. (Or-
iginal)
95. Section of the medulla oblongata at the pyramidal decussation.
305
311
(Original)
96. Nuclei of the cerebral nerves in the medulla, pons, mid-brain,
Motor (or genetic) inter-brain, and olfactory bulb. nuclei red, terminal (or sensory) nuclei blue. (After
Morris's Anatomy) 315 A. Transverse section. (After 97. Meninges of the spinal cord. Key and Retzius.) B. Anterior view. (After Ellis.)
(Morris's Anatomy) 327 Diagrammatic section of the spinal meninges and spinal cord. (After Morris's Anatomy) 329 The arteries and veins in the spinal cord. Diagrammatic. (After Morris's Anatomy) 331 Posterior view of the spinal cord, the dura mater and the arach-
98.
99.
100.
(Brubaker
Sappey)
:
334
A.
The
cervical.
B.
The
thora-
C.
sacral.
(Or-
iginal)
335
of cells, in the cervical
and thora-
Diagrammatic. (In part after Bruce and Cunningham) 341 103. Tracts of fibers and columns of cells in the lumbar and sacral
regions of the cord.
345
(Original). 349
The
Diagrammatic.
105. Direct
to cerebral
and
spi-
367 (Original) 106. Indirect motor paths to the spinal nerves. Diagrammatic.
(Original)
107.
Diagrammatic.
371
tactile,
Common
by way
of the
posterior
column and
Diagrammatic.
(Original)
375
LIST OF ILLUSTRATIONS.
FIG.
XXX111
PAGE
108.
Common
way
379
and spino-thala-
mic
Diagrammatic.
(Original)
(Gordinier after
Van
384
Gehuchten)
optic path.
(Original)
reflex
arc.
387
(Brubaker after Moral and
391
A A
simple
spinal
Dayori)
113.
more complicated
proprii.
392
114.
(Gordinier after E. A.
396
115.
Two
116.
117.
4.25 mm. long. (McMurrich after His) 397 Diagram showing development of neurones in the spinal cord. (McMurrich after Schafer) 398 Ventral and dorsal zones of the spinal cord. (Gordinier and
118.
399
of the third
month.
(Mc406
His)
and one
His) 407
409
(Gordinier
and Quain
after
20.
Diagrammatic
Medial
(Morris's
Anatomy
121.
after
Huxley)
through the brain of an embryo of
fissures
sagittal section
three
on the
411
(McMur-
The
fossa
cerebri
month.
in
123.
Body) Permanent fissures and sulci on the convex surface cerebrum as seen in a seven months' embryo. Murrich after Cunningham]
tricles
413
of the
(Mc414
an embroynic brain, the roof of the lateral venhaving been cut away. Embryo of 12.6 mm.
after
(McMurrich (McMurrich
His)
416
mm.
422
after
His)
XXXI V
FIG.
LIST OF ILLUSTRATIONS.
PAGE
127.
from an embryo of eight weeks. (McMurrich after His) 423 Transverse section of the spinal cord, from an embryo of (A) four and one-half weeks, and (B) of three months.
(McMurrich
128.
after
His)
427
Mode
of origin of anterior
nerves.
and
Diagrammatic.
His)
428
ERRATA.
Page 30.
vesicles.
Paragraph two,
Page 57- Fig- 2 3- "Intraparietal sulcus," and wherever found should be written Interparietal sulcus.
Page
1 08.
Paragraph
word
is
sulcus,
instead of "fissure."
Page 184.
Page 264.
spinal tract,"
Paragraph one,
Fourth
line,
last line
and
first
word
is
Alfred,
and wherever
Ascending an-
Page 265. Paragraph two, third line, the phrase "anterior descending cerebello-spinal tract," should stand Descending anterior cerebello-spinal tract: likewise, wherever the quoted phrase is found.
Third line, last word is (Horsley). Paragraph three, last sentence, the phrase "fasciculus cerebello-spinalis," and wherever else it occurs should stand thus fasciculus cerebello-spinalis (posterior). Page 288. Paragraph three, fifth line, the substantive "cerebello-
spinal fasciculus" should have "posterior" prefixed so as to read posterior cerebello-spinal fasciculus. This is a necessary variation
from the
BNA and
Last
striae.
Page 291.
the medullary
applies wherever the phrase is found. line, there should be added a fifth item, viz., (e)
ERRATA.
XXXV
Fifth line,
common
intermediate nerve.
as
Paragraph three, third item, (3) should be omitted, no posterior root-fibers decussate. Intrinsic fibers only cross through the gray commissure (Mott and Russell).
Page 358.
should be added:
Page 410.
sulcus.
Three membranes
it.
membrane composed
lined
of white fibrous
cells,
and yellow
elastic tissue
its
with
flat
endothelial
it is
which constitute
internal
surface.
which
it
closely adherent to the cranial bones of forms the real periosteum; but it is attached chiefly at
In children
the foramina and along the sutures in adults. The dura of the brain is made up of two layers which are separable up to the
The
bones.
their nutrient
it
membrane.
Through
and sutures
is
continuous with
In the adult the internal layer of the the external periosteum. dura separates from the outer layer only over the apex of the
petrous bone, to form Meckel's space for the semilunar ganglion (Gasseri); at the foramina, to form sheaths for the nerves; and,
Processes.
of the
From
off.
The
falx cerebri
and falx cerebelli hang vertically in the longitudinal fissure of the cerebrum and the posterior notch of the cerebellum; and,'
into the transverse fissure of the cerebrum, extends horizontally the
tentorium
cerebelli.
The
i,
and
4)
is
attached
and behind
tuberance and superior surface of the tentorium; the falx cerebelli (Fig. i) continues from the inferior surface of the tentorium, along the occipital crest, to the posterior border of the foramen
magnum.
(Fig. 2)
is
of
the
tentorium
lateral
cerebelli
and the
arms of
the crucial ridge forward to the petrous bone; and, then, it is along the superior border of the petrous bone to the clinoid processes -of the sphenoid. Between its clinoid attachments there is
tentorii,
The diaphragma
sellae is
dura which covers the hypophyseal fossa. Sinuses. (Sinus dura matrix}. Large venous passages lined with endothelial cells, and called sinuses, are situated between
In the convex and in the layers of the dura (Figs, i, 2, 3 and 4). the free border of the falx cerebri are, respectively, the superior
sagittal sinus
(s.
sagittalis superior)
sinus
(s.
sagittalis
inferior).
The
to the
extends
Having Herophili,) located at the internal occipital protuberance. run through the posterior two-thirds of the concave border of
the falx cerebri, the inferior sagittal sinus joins the great cerebral vein at the margin of the tentorium and forms the straight sinus The latter runs through the middle of the tentorium (s. rectus).
to the confluens
(Fig.
2).
The
occipital sinus
(s.
occipitalis)
magnum upward
the
same
(s.
point.
In
the confluens
rise
sinuses
transversi)
arms
(Fig. 2). Grooving the horizontal each runs outward in the tentorium to
the base of the petrous bone, where it receives the superior petrosal sinus; it then turns downward through the sigmoid fossa, com-
municates with the occipital sinus and unites with the inferior Situated on either side petrosal sinus in the jugular foramen.
the large cavernous sinus (s. cavernosus] (Fig. 3), which receives at the sphenoidal fissure the sinus (s. ala parva], spheno-parietal the course of which is along the posterior border of the lesser
Fig.
i.
showing
d. Great cerea. Falx cerebri. b. Superior sagittal sinus, c. Inferior sagittal sinus, bral vein. e. Straight sinus, f. Tentorium cerebelli. h. Superior g. Transverse sinus, sinus, i. Falx cerebelli. j. Seventh and eighth nerves, k. Ninth, tenth, and petrosal eleventh nerves. 1. Twelfth nerve, m. Second cervical nerve, n. Fourth nerve, o. Third nerve, p. Second nerve, q. Middle meningeal artery, r. Internal carotid artery, s. Vertebral artery, t. Fifth nerve, u. Sixth nerve, v. Inferior petrosal sinus, w. First cervical nerve, x. Ligamentum denticulatum.
cavernous sinus divides into the superior petrosal sinus (s. petrosus superior] and the inferior petrosal sinus (s. petrosus inferior}. The sinus intercavernosus anterior and sinus intercavernosus
posterior extend across the
From
the
bifurcation of the cavernous sinus^at the apex of the petrous bone, the petrosal sinuses run outward along the corresponding superior
and
Fig. 2.
Horizontal section of skull, showing tentorium cerebelli, tentorial notch and sinuses. (After Morris's Anatomy.)
f.
b. Internal carotid artery, c. Optic tract, d. Third nerve, e. Basis Cerebral aqueduct (Sylvii). g. Quadrigeminal body. h. Falx cerebri. cerebelli. j. Straight sinus, k. Crista galli. 1. Optic nerve, m. Spheno-' parietal sinus, n. Middle cerebral artery, o. Anterior cerebral artery, p. Posterior communicating artery, q. Cavernous sinus, r. Superior cerebellar artery, s. Posterior cerebral artery, t. Superior petrosal sinus, u. Free border of tentorium bounding tentorial notch. v. Transverse sinus, w. Superior sagittal sinus. a.
Infundibulum.
pedunculi.
i.
Tentorium
(Figs,
and
2)
the petrous bone; the inferior petrosal sinus, in its course to the jugular foramen, is joined to its fellow, across the basilar process
of the occipital bone,
(p. basilaris]
and, in
the jugular foramen, unites with the transverse sinus in forming the internal jugular vein. Arachnoid Granulations (Fig. 25). Along and within the
Fig.
3.
Sinuses in the base of the cranium, also meningeal arteries. (After Morris's Anatomy.)
a. Meningeal branch of anterior ethmoidal artery, b. Meningeal branch of posterior ethmoidal artery, c. Middle meningeal artery, d. Ophthalmic division of fifth nerve, e. Third nerve, f. Cavernous sinus, g. Fourth nerve, h. Auditory and facial nerves, i. Superior petrosal sinus, j. Inferior petrosal sinus, k. Petro-squamosal sinus. 1. Accessory nerve. m. Sigmoid part of transverse sinus, n. Posterior meningeal branch of vertebral artery. o. Left marginal sinus, p. Left transverse sinus, q. Superior sagittal sinus, r. Circular sinus, s. Carotid artery, t. Sixth nerve, u. Basilar artery, v. Basilar plexus of veins. w. Auditory artery, x. Vertebral artery, y. Glossopharyngeal and vagus nerves, z. Anterior spinal artery, aa. Hypoglossal nerve, bb. Accessory nerve, cc. Right marginal sinus, dd. Occipital sinus, ee. Right transverse sinus.
superior sagittal, the straight, the transverse, the petrosal and the cavernous sinuses are the granulationes arachnoideales (Pacchionian bodies). These granulations are enlarged villi of the
arachnoid
and seem
to
afford
an
They
are said to
make
their
Some of them by absorption produce larger in the male sex. depressions in the cranial bones called joveola granulares.
The
arteries
which
(i)
The
anterior
meningeal from the anterior ethmoidal branch of the ophthalmic. (2) The middle meningeals, viz., the great and the small middle
meningeal from the internal maxillary, the meningeal branch
Fig. 4.
sagittal sinus
and arachnoid
(Gordinier after
Key and
Retzius.)
a.
d.
Dura mater,
Subarachnoid space, b. Superior sagittal sinus, c. Arachnoid granulations (Pacchioni). f. Anterior cerebral artery, e. Falx cerebri. g. Corpus callosum.
and of the internal carotid, and the meningeal branch of the ascending pharyngeal which enters the middle fossa of the cranium through the lacerate foramen. (3) The
of the lacrimal
posterior
meningeal
arteries,
which
rise
pharyngeal, the occipital and the vertebral and are distributed to the dura over the posterior cranial fossa.
is
The great middle meningeal artery (arteria meningea media) much the largest and is most important (Fig. 5). It overlies
Like the other
Ascendarteries
it is
meningeal
it divides near the upper border two large branches, the anterior and posterior. The posterior runs horizontally backward just below the squamoparietal suture and then ascends over the posterior half of the
of the
squamosa
into
parietal
bone.
The
anterior
one inch,
at
and
at
two inches
Fig. 5.
Median
behind the zygomatic process of the frontal bone and above the
zygomatic process of the temporal bone. The following nerves give branches to the dura:
ophthalmic, semilunar ganglion, vagus of the cerebral group; and the sympathetic.
lear,
Troch-
Dura
of the Cord.
and
8
function.
is
In structure
it is
membrane covered
externally with endothelium. Internally it is joined to the pia mater by innumerable fibrous trabeculae, the subarachnoid tissue
(Fig. 4).
The
and
all
subarachnoid
spaces lined with a single layer of endothelial cells, hence both surfaces are formed of endothelium. Conical elevations of
fibrous
villi
constitute
the
Relations.
The arachnoid
dura and
it.
is
prolonged, as a sheath,
pierce
does not dip into the sulci of the cerebrum (Fig. 6); but into the lateral fissure and the longitudinal fissure, and does only not reach to the bottom of the latter. From the pia it is sepaIt
rated by the
cisterna
interpeduncularis,
It is
chiasmatis
and
cistema
located in front of the medulla, pons and mid-brain and between the temporal lobes of the cerebrum.
fossae lateralis cerebri.
The
is
medullaris, a space between the medulla and cerebellum, and the cisterna venae cerebri magnae, situated along the straight sinus and the great cerebral vein. These two, the anterior and posterior,
are the largest subarachnoid spaces and they contain much of But in the cerebral sulci and fissures
The
anterior
subarachnoid
this
space
has
slit-like
com-
membrane, as
merely as a visceral layer of the arachnoid, and regard the endothelial lining of the dura mater as its parietal layer. According to such,
therefore, the subdural space
THE
PIA
Magendi} and the lateral apertures (apertura quarti, Key and Retzii).
laterales ventriculi
The
vessels seen for a short distance in the arachnoid belong Its nerves are doubtful. Perhaps branches
it.
Arachnoid
Third ventricle
Fourth
ventricle
Infundibulum
Cisterna interpeduncularis
Cisterna cerebello-medullaris
Cisterna pontis
Fig. 6.
Diagram
tum
it
fewer trabeculae join it to the pia; are collected to form a fenestrated seppart, hi the posterior median line (Fig. 97, A). The external
of the cord
by the
lie
between
and the
pia.
It
is
a vascular
membrane com-
10
outer surface
trabeculae.
The endothelium covering its continuous with that ensheathing the subarachnoid The pia closely follows the brain surface (Fig. 6).
is
Internally,
it
transmit
blood-vessels;
sends supporting trabeculae into the brain, which and externally it forms an investing
Two
The chorioid
important processes are formed by the pia mater tela of the third ventricle (tela chorioidea
:
ventriculi tertii) is
pushed forward into the anterior part of the transverse fissure of the cerebrum between the fornix and the
interbrain (Fig. 6). Hence the old name, velum interpositum. It is triangular in shape, with apex directed forward (Fig. 7). Each lateral border is tucked into the chorioidal fissure of the cerebral
the median part of the fold is in the roof of the third ventricle. Between the two layers of this chorioid tela is some areolar tissue
through which run backward the two internal cerebral veins and unite near the base of the tela to form the great cerebral vein.
The
ventricles oc-
cupy, respectively, the lateral borders and the median area of this chorioid tela. second fold of pia mater is tucked .into (2)
oblongata and ventral to the posterior median part of the cerebellum (Fig. 6). It is called the chorioid tela of the fourth
ventricle
(tela
chorioidea ventriculi quarti) because its inferior and contains the chorioid plexus of that
medulla and the roof -epithelium of the fourth ventricle (Fig. 8). It is pierced by three foramina which are situated as follows:
One
over each lateral angle of the fourth ventricle, the lateral The apertures (Key and Retzii), and one over its inferior angle.
latter is the largest
and is called the median aperture (Magendi). Those three foramina establish communication between the posterior subarachnoid space and the fourth ventricle.
The
11
and
arteries of the pia mater supply the brain (Figs. 9,- 10, 12). They are the anterior, middle and posterior cerebrals;
the anterior and posterior chorioidals; and the anterior and pos-
THE
PI A
II
Fig.
7.
and
THE
PIA
13
many
branches.
The veins
the internal
are
and great cerebral veins, the veins of the chorioid plexuses of the lateral, third and fourth ventricles and the basilar vein; the cerebral veins, superior, medial and inferior; and the
superior and inferior cerebellar veins. the sinuses (see page 2).
All of
them empty
into
Frenulum
Lateral
veli
Fourth nerve
fillet
Superior medullary
velum
Fourth
ventricle
Restiform body
Taenia
Epithelial roof of fourth ventricle Cuneate tubercle
Inferior medullary
velum
Chorioid plexus
Clava
Median aperture
(Magendi)
Tuberculum
cinereurr*
Obex
Fig. 8.
Roof and
and
its
chorioid plexuses.
and nth
and the sympathetic supply the pia mater and its blood-vessels. The pia mater of the spinal cord has two layers, the outer of which is the more vascular and contains the spinal arteries and
the tributaries of the external spinal veins. It forms three processes, namely, the anterior septum, which occupies the anterior median fissure, and the ligamentum denticulatum of each side.
14
The
brain
is
(a,
and the posterior communicating; vertebralis) gives off the anterior and
inferior cerebellar
and then
and forms
The
basilar artery
(a. basilaris]
runs upward
and terminates
in the
the anterior inferior cerebellar, the pontal, the internal audiCertain of these arteries form tory and the superior cerebellar. a wonderful circular anastomosis at the base of the brain, called
viz.,
is
to the cerecere-
brum
(Figs. 9
and
rhombencephalon (pons,
The
cephalon.
A.
The
ies
Arterial Circle.
The
arter-
which supply the cerebrum freely communicate in the arterial circle, which is really a heptagon extending from a point in the
longitudinal fissure anterior to the optic chiasma, back to the pons It is about an inch and a half long, and from a half (Fig. 9).
to
cerebral arteries converging forward from the internal carotids and, through the anterior communicating artery (a. communicans
anterior], uniting just as they enter the longitudinal fissure of the
cerebrum.
These
vessels
form three
sides of the
heptagon and
On
communic-
ating artery (a. posterior] carotid with the posterior cerebral artery, forms the lateral bound-
communicans
15
The
posterior
cerebral
arteries
bound
the
behind, and so complete it (Fig. 10). The large distal branches of the arteries which are connected with the arterial
circle are distributed
hemispheres;
while
the
Fig. 9.
Arterial circle
and its branches on the base of the brain. (After Morris's Anatomy.}
d. g.
terior inferior cerebellar artery, h. Vertebral artery, i. Posterior inferior cerebellar artery. Anterior communicating artery, k. Antero-lateral ganglionic. 1. Anterior chorioid. j. m. Posterior communicating artery, n. Posterior chorioid. o. Basilar artery, p. Hemisphere of cerebellum cut away. q. Anterior spinal artery.
c. Internal carotid artery, b. Middle cerebral artery, a. Anterior cerebral artery, terp-median ganglionic. e. Posterior cerebral artery, f. Superior cerebellar artery,
Pos-
An-
ganglia and the inter-brain. The former belong to the cortical system (Ai), the latter to the ganglionic system (A2). Al. The Cortical System of Arteries. The cortical arterial
system comprises the distal portions of the anterior, middle and The branches of posterior cerebral and the chorioidal arteries.
16
these great vessels pierce the hemispheres perpendicularly to the surface. They are distributed, the short, to the cortex, and the
long, to the
To
a limited extent
(a. cerebri
anterior, Figs. 9
and
12), a branch of the internal carotid, runs forward and toward the median line above the optic nerve and enters the longitudinal
fissure;
it
is
here joined to
anterior communicating.
mate by a very short artery, the Winding around the genu of the corpus
its
callosum,
it
runs back on the medial surface of the hemisphere to It gives origin to the antero-median
The
and
the medial orbital gyrus, the gyrus rectus, the olfactory bulb, tract, medial and intermediate
triangle,
and parolfactory area. (2) The anterior internal frontal (Fig. 12) which enters the anterior parts of the gyrus cinguli and superior frontal gyrus on the medial surface and of the superior and middle frontal gyri on the convex surface. (3) The middle internal frontal
branches, which are distributed to the middle part of the gyrus cinguli, to the paracentral lobule and to the upper portions of the
and
(4)
The
to the
occipito-parietal sulcus.
They supply nearly the whole corpus the posterior half of the gyrus cinguli, a part of the callosum, paracentral lobule, the praecuneus, and the superior parietal
lobule.
The middle
of the
it
cerebral artery
(a. cerebri
n)
lateral fissure
cerebrum
where
breaks up into several parieto-temporal branches. It gives origin to the antero-lateral ganglionic arteries, and to four cortical
branches:
(i)
The
lateral
orbital
the anterior and posterior orbital and the inferior frontal gyri. (2) The ascending frontal, two branches, which follow the precentral
and central
sulci,
17
the posterior fourth of the middle frontal gyrus. (3) The ascending parietal, whose course is along the interparietal sulcus, furnishes
parts
blood
of
to
and inferior parietal lobules. (4) The arteries, which comprise two polar branches to the temporal lobe and a large posterior branch. The latter runs in the posterior ramus of the lateral cerebral fissure to its upturned posterior end and there bifurcates into a parietal and a temporal branch, which just pass the anterior limit of the occipthe superior
ital lobe.
is
The
to the temporal pole and to the superior, middle and part of the inferior temporal gyri; to the major parts of the supramarginal, angular and post-parietal gyri, and to a very small portion of the superior
and
cerebri
posterior},
ter-
boundary
of the
joined to the internal carotid by the posterior communicating artery (Figs. 10 and 12). It winds backward between the midb^ain and gyrus hippocampi to the tentorial
is
and
surface of the cerebral hemisphere where, just beyond the splenium of the corpus callosum, it terminates in the calcarine and occipito-
From the posterior cerebral arteries originate parietal branches. the postero-median and the postero-lateral ganglionic, and two or more posterior chorioidal arteries and three cortical
branches: (i) The temporal branches, often an anterior, middle and posterior temporal, which supply the hippocampal and the
fusiform gyri and a part of the lingual and of the inferior temporal gyrus. (2) The calcarine artery, which runs along the fissure of the same name and supplies the cuneate and lingual gyri; also the pole and the lateral
and superior
(3)
occipito- parietal artery, a single branch, which runs along the sulcus occipito-parietalis over the supero-medial border to
The
the convex surface of the cerebral hemisphere and is distributed to the cuneus, the praecuneus and the superior occipital gyrus.
The
posterior
chorioidal
and
18
the posterior cerebral which run forward in the transverse and chorioidal fissures of the cerebrum to the chorioid plexuses of the lateral and third ventricles (Fig. 7).
The
(a.
from the internal carotid artery just proximal to its anterior and middle cerebral branches, and runs backward and outward along
the optic tract to the anterior inferior end of the chorioidal fissure, which it enters (Fig. 10). It terminates in the chorioid plexus of the inferior horn of the lateral ventricle, and gives collateral
tract,
dentata, the hippocampus, the crus of the fornix part of the internal capsule.
from the
circle
A2. The Ganglionic System of Arteries. Small arteries arterial circle and from the cerebral arteries near the
constitute
this
their distribution
system (Fig. 10). The arteries pass to without communicating with one another or
They are the end-arteries of Cohnheim. and ganglionic systems, there is an area poorly supplied with blood. That is the area of cerebral softening in old age. The ganglionic system of arteries is made up of six of small vessels: The antero-median, the right and left groups antero-lateral, the postero-median and the right and left posterowith the cortical arteries.
cortical
Between the
lateral.
cerebrals in front of the optic chiasma (Fig. 10). They supply the chiasma, the lamina terminalis, the rostrum of the corpus
callosum, the septum pellucidum and the head of the caudate nucleus.
The
arterial
either side,
antero-lateral ganglionic arteries take their origin, on from the middle cerebral artery, a little outside the
circle (Fig. 10). They pierce the anterior perforated substance and are distributed to the striated body, internal capsule and thalamus. The largest one of this group is the lenticulostriate artery.
It supplies
its
On account
of
the greater part of the corpus striatum. frequent rupture, it is called the artery of cerebral
hemorrhage (Charcot).
Fig. 10.
Arterial circle
and its branches on the base of the cerebrum. (Gordinier after Duret.)
On the left side of the brain the temporal lobe is cut away so as to open the inferior and posterior horns of the lateral ventricle. The mid-brain is divided close above the pons and the posterior cerebral arteries are cut at their origin from the basilar. Ganglionic arteries: am. Antero-median group arising from the anterior cerebral, al. Antero-lateral group, from the middle cerebral, pm, pi (on the optic thalamus). Posteromedian and postero-lateral groups, from the posterior cerebral. Chorioidal arteries: a ch. Anterior, from the internal carotid, p ch (on the splenium). Posterior, from the posterior cerebral. Cortical arteries: I, I. Inferior internal frontal, from the anterior cerebral. 2. Inferior external frontal. 3. Ascending frontal. 4. Ascending parietal, and 5, temporo-parietal from the middle cerebral. 6. Anterior temporal, 7, posterior temporal, and 8, occipital, from the posterior cerebral.
21
arteries
They supply the interpcduncular structures, (Figs. 9 and 10). the peduncles and, after piercing the posterior perforated substance, the walls of the third ventricle and the medial parts of
the thalami.
They
rise,
on
either
from the posterior cerebral artery after it has wound around the base of the peduncle (Fig. 10). They are distributed to the of the thalamus; the geniculate, quadrigeminal and posterior part
pineal bodies; the quadrigeminal brachia and the pedunculus The superior cerebellar arteries send several branches cerebri.
to the
dorsum
of the mid-brain,
arterial
supply
of the cerebrum.
mate
in forming the
The
(v. cerebri
interna)
is
formed by
the union of the chorioidal, the terminal and the vein of the septum pellucidum. It runs backward between the layers of the
chorioid tela of the third ventricle
(Fig.
7),
receiving several
tela,
bodies and the corpus callosum and, finally, it receives the basilar vein from the inferior surface of the cerebral hemisphere. Under
the splenium of the corpus callosum it joins the internal cerebral vein of the opposite side and forms the great cerebral vein.
(v. cerebri
it is joined by the inferior sagittal and then continued as the sinus rectus. This short vein receives collateral tributaries from the gyrus cinguli, from the
22
medial and tentorial surfaces of the occipital lobe and from the superior surface of the cerebellum (Cunningham).
all
and form the external veins. The External Veins of the Cerebrum. The external cerebral veins (vena cerebri externa) are numerous and of large size. They
ramify in the pia mater and in the subarachnoid space. They empty into the dural sinuses, as a rule, against the current in the
sinuses,
and
they
form
The
superior
Fig.
1 1
Middle cerebral artery and branches. (Gordinier after Quain and Charcot.)
Antero-lateral group of ganglionic arteries. I. Inferior external frontal artery. 2. Ascending frontal artery. 4. Parieto-temporal artery. 3. Ascending parietal artery.
CENT.
cerebral
and the
inferior cerebral,
the
medial
The superior cerebral veins (vena cerebri superiores), twelve or more in number, carry away the blood from the superior surface of the hemisphere. They run obliquely upward and forward
into the superior sagittal sinus.
(Vena
cerebri mediales}.
They
23
into the superior
The
empty
and the
much
inferior cerebral veins (vena cerebri inferiores) drain the base of the cerebrum and the lower border of its convex surface.
The
the tentorial surface of the hemisphere, from three to five of these veins empty into the transverse and superior petrosal sinuses. Those from the temporal and frontal lobes empty into
On
the
spheno-parietal sinus and cavernous sinus, excepting the small anterior cerebral vein and the deep middle cerebral vein,
accompanies the artery of the drains the gyrus cinguli and corpus callosum, chiefly; and, in the fossa lateralis cerebri, unites with vessels that descend from the corpus striatum, the inferior striate veins, and
The
same name.
with the deep middle cerebral vein. The deep vena cerebri media drains the insula and the opercula, in part, and deep in the fissure runs medianward to the fossa lateralis cerebri and helps to form The basilar vein (v. basilaris), is formed at the the basilar.
anterior perforated spot by the deep middle cerebral, the inferior striate and the anterior cerebral veins. Running backward it
receives
the hippocampal gyrus and the inferior horn of the lateral ventricle, and from the mid-brain, as it winds around it to empty into the
corresponding internal cerebral vein near its termination. In the fissura lateralis cerebri (Sylvii) runs also a superficial vein, called
the superficial middle cerebral (v. cerebri media) which receives tributaries from the surfaces adjacent to the posterior ramus and the stem of that fissure and empties into the cavernous sinus; but
it
other outlets,
viz.,
The connection occasionally established superior sagittal sinus. between the superficial middle cerebral vein and the transverse
sinus
is
produced when
it
joins
one of the
superior cerebral veins. great anastomotic vein connects the superior sagittal with the cavernous sinus.
The
24
cord; perivascular
to the
subarachnoid spaces.
B.
Bl. The medulla oblongata is supplied with blood by the following branches of the vertebral artery: The posterior and the anterior spinal, the posterior inferior cerebellar and several
short bulbar arteries (Fig. 9).
(a. cerebelli inferior posterior}
The
the medulla, runs between the vagus and accessory nerves, enters the vallecula cerebelli and gives branches to the medulla and to
The
anterior spinal
formed by the
artery,
and the
medulla, descends in front of the posterior lateral sulcus. Both The branches for the distribute branches along their course.
nerves, suggesting the centrifugal
most part enter the median raphe or follow the roots of the bulbar and centripetal arteries of the
spinal cord (Fig. 13).
the
same course
as the arteries.
of the
The
is
pons and
joins the ventral veins drained into the cerebellar veins or directly
median vein
The
posterior
median vein
bifur-
and the two branches wind around the medulla to its anterior surface and empty into the inferior petrosal sinus or the basilar plexus. Issuing from
cates y-like at the middle of the medulla
the medulla with the roots of the ninth to the twelfth cerebral
nerves are three or four small veins, the radicular veins, which run into the occipital and inferior petrosal sinuses (Cunningham).
Both arteries and veins possess perivascular lymph spaces, but there are in the medulla no lymphatic vessels. B2. The pons Varolii is supplied by the pontal, the superior cerebellar
artery (Fig.
and the posterior cerebral branches of the The short and transverse branches of the 9).
basilar
basilar
artery, the pontal arteries (aa. pontales), furnish the greater portion
of blood to the basilar area of the pons, while the superior cere-
27
artery
supplies
the
superior
brachia conjunctiva cerebelli. The branches enter the median raphe, also the substance of the pons elsewhere, especially along
the nerve roots, and run at right angles to the surface into The deep veins of the pons run forward and form a plexus on
it.
its
surface which, according to Cunningham, is drained by a superior efferent into the basilar vein and by an inferior efferent into the cerebellar veins or the superior petrosal sinus. There are no
Fig. 13.
a. spin. post.
Vertebral artery,
artery.
lymphatic vessels in the pons; but, as elsewhere in the central nervous system, there are lymph spaces about the blood-vessels.
is
furnished by
The superior cerebellar, from three pairs of arteries (Fig. 9). the basilar, supplies all the superior surface except a narrow zone at the posterior border; the anterior inferior cerebellar, also from
the basilar, and the posterior inferior cerebellar, from the vertebral, supply the inferior surface and the posterior part of the
superior surface.
The Superior Cerebellar Artery. (A. cerebelli superior}. Rising from the basilar just behind the posterior cerebral, from
28
which
is
it
winds dorsally
it
where
bifurcates
and a
lateral
branch
(Fig. 9).
The
medial branch
continues along the trochlear nerve in the groove between the cerebellum and the mid-brain almost to the median line and then,
;
worm
of the cere-
posterior
extremity.
It
distributes
branches to
chorioidea
geniculate
bodies,
corpora
quadrigemina,
tela
Fig. 14.
Median
section of
(McMurrich
I.
Myelencephalon. II. Metencephalon: i, Pons, 2, Cerebellum, in. Isthmus rhombencephali. IV. Mesencephalon: i, Pedunculi, 2, Corpora quadrigemina. V. Diencephalon: I, Pars mammillaris hypothalami, 2, Thalamus, 3, Epithalamus. VI. Telencephalon: I, Pars optica hypothalami, 2, Corpus striatum, 3, Rhinencephalon, 4, Neopallium.
ventriculi
tertii
of
vermis superior cerebelli and the medial part of the superior surface of the hemisphere. The lateral branch of the superior
point of origin near the sulcus lateralis of the mid-brain onto the superior surface of the cerecerebellar artery passes
its
from
bellum.
It
its course. The lateral branch, with the medial, supplies the superior cerebellar surface together almost as far back as the horizontal sulcus of the cerebellum,
2Q
anterior
The anterior
its
(a. cerebelli
inferior, Fig. 9) is
inferior
and middle
(Sometimes
runs
vessels.)
it
is
In
its
keeping close to the anterior border of the hemisphere. course it passes anterior to the abducent nerve and posterior
to the facial
It supplies
of the
posterior inferior cerebellar artery (a. cerebelli inferior posterior, Fig. 9) is the largest branch of the vertebral and is given off just before the vertebral arteries unite and form the basilar.
The
Passing first between the root-bundles of the hypoglossal nerve and then between those of the accessory and vagus nerves, the
posterior inferior cerebellar artery bends at a right angle back-
ward and runs between the medulla and the cerebellar hemisphere, where it divides into a medial and a lateral branch. The medial branch follows the sulcus valleculae and gives branches to the medial part of the hemisphere and the vermis inferior. It anastomoses with
fellow of the opposite side. The lateral branch, runs lateralward from the posterior cerebellar notch over the
its
branches wind
around the postero-lateral border and communicate with the superior cerebellar artery on the upper surface of the hemisphere.
The undivided trunk of the posterior inferior cerebellar artery gives small branches to the medulla oblongata and supplies the chorioid tela of the fourth ventricle. The
internal cerebellar veins bring the blood from the interior
of the organ
veins.
cerebellar veins (vena cerebelli supeconverge forward into a medial vein, which empties into
the great cerebral vein, and several lateral veins, which end in the transverse or the superior petrosal sinus.
The
riores] also
vein,
and upward
and a
number
of lateral veins.
The
lateral
in the cere-
bellum, but the perivascular lymph spaces carry out the lymph and pour it chiefly into the subarachnoid space.
TABLE
In accordance with
I.
its development the brain or encephalon divided into three embryologic divisions which comnaturally prise the derivatives of the anterior, the middle and' the poste-
Cerebral Hemispheres
Corpus Callosum
Fornix
Anterior Commissure
Septum Pellucidum
i.
End-brain, or
Telencephalon
Lateral Ventricles
I.
Fore -Brain, or
Prosencephalon
(Ant. vesicle)
-,
Foramina
ularia
interventric-
Aula of
third Ventricle.
Thalami
Corpora Mammillaria
2.
Inter -brain, or
Corpus Pineale
Corpora Geniculata Third Ventricle, excepting the aula.
Diencephalon
Cerebrum
<
Bases Pedunculi
Pedunculi
Cerebri
II.
Substantia Nigra
Mid-Brain, or
(of
Lamina
Quadrigemina
Corpora
ina.
Quadrigem-
Brachia.
Prosencephalon. (forebrain)
Cerebrum
j-
Mesencephalon
(mid-braiu)
Encephalon
(brain)
--Cere-
1
!
bellum
Metencephalon
(hind-brain)
\
Rhombencephalon
(Medulla oblongata)
'
Pars cervicalis
")
'
Pars thoracalis
- Pars lumbalis
Fig. 15.
Diagrammatic.
33
Metencephalon
(Hind-brain)
Upper
half of
Fourth Ven-
III.
Rhombencephalon
(Post, vesicle)
tricle
Myelencephalon
(After-brain)
The cerebrum embraces the fore-brain and the mid-brain, as shown by the table. So we may make a more comprehensive
division of the brain into only
two grand
:
divisions:
The
great
brain or cerebrum and the rhombencephalon (Fig. 15). now simplify the above table as follows
I.
We may
Cerebrum, embracing
End-brain, or Cerebral Hemispheres,
Inter-brain
etc.
Mid-brain.
II.
Pons
Medulla Oblongata.
CHAPTER
II.
do
brain forms the greatly expanded superior extremIt is derived from three sack-like ity of the cerebro-spinal axis. dilatations of the epiblastic neural tube, called the anterior, the
The human
(Fig. 16).
The
cavities of
median
series extend-
ing from the canal of the spinal cord up to the level of the cerebral hemispheres; at that level the central cavity bifurcates into a branch for each hemisphere of the cerebrum (Figs. 17 and
1 8).
Thus
is
formed the
hemis-
aqueduct in the mid-brain, and the fourth ventricle in the hindbrain and after-brain. The walls of these simple embryonic
cavities
undergo
wonderful
development
ultimately they produce all the multiform tures of the adult human brain.
Superior View. The superior surface of the brain is markedly convex (Figs. 19 and 22). It is elliptical in outline, the major
axis being contained in the axis
is
median
is
situated a
little
points which,
parietalia.
when
the brain
This surface
is closely
34
35
calvaria. Only the great convoluted hemispheres of the cerebrum are visible from the superior view-point. The two hemispheres are separated by a deep, median cleft, called the longitudinal
fissure
the cerebrum (fissura longitudinalis cerebri) from which the falx cerebri has been removed.
of
Posterior View.
great structures
When
the brain
is
Fig. 1 6. Diagrams of surface views and sections of germinal areas showing the development of the primitive streak, neural groove, neural tube and brain vesicles. (After Morris's Anatomy.)
A. Earlier stage, a. Germinal area. b. Neural groove, c. Primitive streak. B. .Later stage, a. Germinal area. b. Fore-brain (rudiment of cerebral hemispheres), c. Optic vesicle. d. First cerebral vesicle, e. Second cerebral vesicle, f . Third cerebral vesicle, g. Primitive streak. A'. Section through area along the line a. a. Germinal area. b. Neural groove. B'. Section through area along line b. a. Germinal area. b. Neural crest, c. Neural tube.
their
irregular gyri
and sulci; second, the transversely laminated cerebellum, lying below the cerebrum and separated from it by the transverse fissure of the cerebrum (fissura transversa cerebri); and third, the inferior extremity of a relatively small median
structure, the
medulla oblongata.
The cerebellum
is
especially
30
characterized by
fied
appearance. partial hemispheres produced by a posterior median depression, called the posterior cerebellar notch, and by a longitudinal groove on its
inferior surface,
fitted
is,
shows a
The
vallecula' is
over the posterior surface of the medulla. The cerebellum therefore, separated from the medulla oblongata by a sharply
Fig. 17.
Diagrammatic horizontal
(Morris's
Anatomy
after Huxley.')
a.
j.
tricle,
Metencephalon. b. Thalamus. c. Medulla oblongata. d. Cerebellum, e. Lateral venf. Olfactory diverticulum. g. Lamina terminalis. h. Corpus striatum. i. Mid-brain. Pineal body. k. Interventricular foramen.
curved, rainbow-shaped fissure. That fissure is the transverse fissure of the cerebellum (fissura Iransversa cerebelli) which, as already pointed out, is bridged over by the arachnoid and contains the cisterna cerebello-medullaris.
of the brain presents three areas, situated in three successive levels, which correspond in location
Inferior View.
The base
fossae in the
occupies the highest level; the middle area is intermediate in position; it occupies the middle fossa and together with the anterior area comprises all of the base of the cerebrum which is visible in
the complete brain; and, the posterior area, which is but the base of the rhombencephalon, is situated at the lowest level in the
posterior fossa of the cranium. The anterior area of the base of the brain
lateral halves
is
divided into
by the longitudinal
fissure of the
cerebrum, and
37
separated from the middle area by the fossa and fissura lateralis The frontal lobe of the cerebral hemisphere, on either cerebri.
side of the longitudinal fissure,
The
is
to the
adapted convex orbital plate of the frontal bone; its medial border most prominent and presents, near the longitudinal fissure, an
(if it
this area.
is
Fig. 18.
Diagrammatic
(Morris's
Anatomy
a. Corpora quadrigemina. b. Mid-brain, c. Pineal body. d. Cerebellum (hind -brain). Medulla oblongata (after-brain), f. Pons Varolii (hind-brain), g. Lateral ventricle. h. Cerebral hemisphere, i. Corpus striatum. diverticulum. k. Pedunculi j. Olfactory cerebri. 1. Thalamus, m. Inter-brain, n. Hypophysis. O. Interventicular foramen. 4. Fourth ventricle. 5. Aqueduct of cerebrum. 3. Third ventricle.
e.
and a white strand, the olfactory tract. Running backward from the bulb, parallel with the longitudinal fissure of the cereoff)
brum
bifurcate into
two
distinct striae,
a medial and a
lateral.
is
prom-
where it is formed by the temporal lobes of It is depressed in its median portion and thus
adapted to the hypophyseal region of the cranial floor. This the end of the longitudinal fissure, in front, backward to a great white, transversely
it contains several important bases pedunculi; posterior perforated substance; the mammillary bodies; tuber cinereum and stem of the infundibulum; optic chiasma, tracts and nerves; lamina cinerea
terminalis;
and the anterior perforated substance. Issuing from the under surface of the cerebral hemisphere and
38
running downward toward the median line, there may be seen a white striated band, a half-inch broad, called the basis pedunculi,
into
which, on approximating its fellow in the median plane disappears the pons. Anteriorly, the X-like optic chiasma (chiasma
opticum) is easily identified near the longitudinal fissure; its anterior limbs are the optic nerves and its posterior, the optic tracts The optic tract, when traced backward and out(Fig. 21).
is
observed to cross
the basis pedunculi at its point of emergence from the cerebral hemisphere. Thus the optic tract and the basis pedunculi form the lateral boundary of a diamond-shaped space extending from
the optic chiasma, in front, backward to the pons. This is commonly called the interpeduncular space. You observe in it three
(i) gray eminence just behind the optic chiasma called the tuber cinereum; (2) a pair of white, nipple-like bodies,
structures:
an eighth of an inch in diameter, known as the white or mammillary bodies (corpora mammillaria), and (3) a triangular, perforated
mass
the infundibulum projects downward and forward from the center of the tuber cinereum and connects it with the hypophysis cerebri;
but
left
usually broken in removing the brain and the hypophysis behind in the hypophyseal fossa. If the optic chiasma be drawn slightly downward and backit is
ward, a transverse and nearly vertical sheet of gray matter will be seen extending upward from it, between the cerebral hemisThat is the lamina cinerea pheres, toward the corpus callosum.
terminalis.
It
bounds
Lateral to the optic chiasma and anterior to the optic tract, the gray substance is perforated by many vessels; it is called the anterior perforated substance
udinal fissure of the cerebrum.
to distinguish it from a similar located between the bases pedunculi. posterior region Posterior Area. The posterior area of the base of the brain
is
formed by the pons, the cerebellum, and the medulla oblongata, which constitute the rhombencephalon (Fig. 21). The pons and medulla are median structures. They are separated by a
39
Fig. 19.
(Original.)
41
the
marked transverse groove containing the roots of the sixth, seventh, the intermediate and the eighth cerebral nerves.
transverse strands of the pons traced lateralward are observed to form a large round bundle, called the brachium pontis, which
The
Be-
tween those pontine strands, at the lateral border of the pons, there should be noticed the roots of the great trigeminal nerve. A sagittal line through this nerve at its attachment to the pons
may be
bellar hemisphere.
regarded as the boundary between the pons and the cereThe hemispheres of the cerebellum form the
appearance
is
the medulla oblongata. The medulla is about an inch long and three-quarters of an inch broad near the pons, but measures less than one-half inch in
already familiar.
is
It is partially
by the anterior median fissure, which is deep, above, but is almost obliterated in the lower half of the medulla by the crossing of the
lateral
pyramidal
tracts,
median
The olive, and the restiform body. bounds the anterior median fissure. It is pyramid (pyramis] an eighth of an inch in width, is most prominent near the pons and tapers off inferiorly because about 80 per cent, of its fibers
pyramid, the
cross over to the opposite side
It is
bounded
and from a
flat
medulla.
surface of
The
the medulla;
The
nent,
olive is equal in
is
promi-
in outline.
The
posterior
lateral
sulcus (sulcus lateralis posterior) separates it from the restiform body. The roots of the ninth, tenth and eleventh cerebral nerves, which are contained in that groove and the restiform
body which
lies
beyond
it,
42
The Roots
of the
(Fig. 21).
TJie
cerebral nerves (nervi cerebrales) are numbered from before backward according to the order of their points of attachment to the
brain surface.
roots, points of
Those points of attachment are, for the motor exit from the brain (apparent origins); and are
The
the real origin of each motor root, and the terminal nucleus (nucleus terminalis], which contains the real central ter-
which
is
mination of every sensory root, are imbedded within the brain substance and do not at present concern us.
1. The olfactory nerves (nervi olfactorii) are the first. .They are the nerves of smell. They are composed of twenty or thirty
which
rise
from the
olfactory cells in the nasal mucous membrane and, passing through the cribriform plate of the ethmoid bone, enter the under surface
of the olfactory bulb.
their
central
The
is,
therefore,
apparent proceed some distance into the gray substance of the olfactory bulb, which constitutes the terminal nucleus of the first nerves, and there
termination.
fibers
The
branch richly and end in relation with the mitral and bush-cells
(real central termination).
2.
Optic Nerve.
is
(Nervus
options).
The second
It rises in
nerve, the
nerve of sight,
its
fibers are
imbedded hi
neuroglia.
the eyeball and into the optic chiasma where the "nerve"
fibers
Passing through the chorioid and sclera of the optic foramen of the sphenoid bone, it enters
is
and the midwhose surfaces they pierce (apparent central termination); brain, they end (real central termination) in the lateral geniculate body,
optic tracts
and
colliculus of
(n.
nerve to the eye (Fig. 21). It issues from the mid-brain at the medial border of the basis pedunculi, which is its apparent origin,
43
Fig. 20.
(Original.)
45
a mass of gray substance, the genetic nucleus (n. originis], situated within the depths of the mid-brain. 4. Trochlear Nerve. The fourth is a motor (AT", trochlearis}.
real origin is in
nerve to the eye and is the smallest of the cerebral nerves. It may be seen winding forward over the basis pedunculi (Fig. 21). Its apparent origin is from the dorsal surface of the brain stem
at the junction of the
;
Fig. 44) this apparent origin cannot be seen in the complete brain. The genetic nucleus of the fourth nerve is located below that of the third in the mid-brain.
is
Trigeminal Nerve. (N. trigeminus). The trigeminal nerve a mixed nerve, motor and sensory (Fig. 21). It is attached to the ventral surface of the pons a little above the middle of its
5.
lateral
border.
The
small anterior
;
apparent central termination of the large sensory root, which rises in the semilunar gangthis point (apparent origin) this is the
but
lion (Gasseri)
close to the
emergence of the
motor
6.
root.
abducens]
is
a motor nerve to
the eye. from the pons at its inferior border, or from the transverse groove between the pons and the medulla, just above the pyramid of the medulla and nearly in line with the
anterior lateral sulcus (Fig. 21).
In the transverse groove between the pons and the medulla, lateralward from the root of the sixth nerve, are the roots of the
seventh,
The
seventh
is
smaller in
diameter than the eighth and medial to it in position; the intermediate is between these two (Figs. 21 and 45). 7. The facial nerve (n. facialis} is the motor nerve to the muscles of expression (Figs. 21 and 45).
in the pons,
its
medulla and pons constitutes its apparent origin. The intermediate nerve (n. intermedius) is so closely associated with the
facial nerve that
many
regard
is
it
in reality a
efferent
whose function
is
taste.
and trophic functions and afferent The efferent fibers rise from the
46
salivary nucleus
pons.
They
issue
facial
between the
from the transverse ponto-medullary groove and auditory nerves (apparent origin) at the
The sensory point where the afferent fibers enter the brain. part of the intermediate nerve, which is the nerve of taste to the
anterior part of the tongue, takes its origin in the ganglion geniculi situated within the canalis facialis (Follopii); its apparent central
termination
8.
is
the
The acustic nerve (n. acusticus] is a sensory nerve, having double function of hearing and equilibrium (Figs. 21, 44
45).
It rises
and
from the
spiral
situated
and
its
bottom of the transverse groove separating the pons from the medulla. The roots of both the seventh and eighth nerves are
near the upper end of the posterior lateral sulcus of the medulla
oblongata.
Glossopharngeal Nerve. (N. glossopharyngeus}. This is a complex mixed nerve, containing efferent -fibers (motor, vasodilator, secretory and trophic) and afferent fibers, which are both
9.
common sensory and gustatory. It is joined to the medulla in the bottom of the superior end of the posterior lateral sulcus This point of attachment is the apparent (Figs. 21 and 45).
origin of
its
efferent
central termination of
its
afferent fibers.
The
and petrosal
glossopharyngeal ganglia situated in the jugular foramen. The genetic nucleus of the efferent fibers is located inside the medulla.
tenth and eleventh nerves.
Behind the ninth nerve in the same groove are the roots of the The roots of the ninth and tenth
are situated between the olive and the restiform body; but, if the nerve trunks have been cut, it is impossible to determine which of the ten or a dozen root bundles belong to each of them. 10. The Vagus Nerve. (N. vagus}. The efferent -fibers of the vagus, like the glossopharyngeal, take their apparent origin from the posterior lateral sulcus, and in the same sulcus the afferent
fibers
and
enter the medulla, apparent central termination (Figs. 21 Its efferent fibers comprise It is a very complex nerve. 45).
47
Fig. 21.
a.
Base of brain.
(Original.)
Olfactory bulb. b. Olfactory tract, c. Medial and lateral olfactory strise. d. Trigonum olf actorium. e. Area parolf actoria (Brocas). f. Anterior perforated substance, g. Optic chiasma. h. Optic tract, i. Tuber cinereum. j. Infundibulum. k. Hypophysis. 1. Corpus mammillare. m. Posterior perforated substance, n. Basis pedunculi. o. Sulcus parclfactorius anterior. 2 to 12, the cerebral nerves.
49
The
jugulare and
g.
and nodular ganglia of the nerve (g. nodosurn) within and just below the jugular fora-
men.
fibers.
11.
(n. accessorius)
is
composed
of a
cerebral
(Fig.
of
which are
efferent in function
The
below the
of the olive
and immediately
This
is
distributed entirely
by way
of
The spinal root (radix spinalis), having taken its apparent origin from the lateral surface of the spinal cord and passed through the foramen magnum, joins the cerebral (acces12.
Hypoglossal Nerve. (N. hypoglossus}. The twelfth is the great motor nerve to the tongue (Figs. 21 and 45). A half
dozen or more radicals make
issue in linear series
it
up; they
rise hi
medulla
between the pyramid and the olive (apparent origin). The root bundles which emerge from the same sulcus below the level of
the olive belong to the anterior root of the first cervical nerve. The student should now turn back to Table I. Study it care-
and identify all the primary and secondary divisions of the brain (Figs. 14, 15, 16, 17, 18, and 28).
fully
CHAPTER
III.
THE CEREBRUM.
The cerebrum with its great hemispheres is that part of the brain which especially characterizes man. In man only do the hemispheres reach such predominant development. Though
they are mere outgrowths of the anterior brain-vesicle in the beginning, they completely overshadow all other parts of the
brain by the seventh month of embryonic life, extending farther forward, backward and lateralward than any other part. Within the cerebral hemispheres lies the physical basis of all mental function; they constitute the central mechanism of thought and consciousness.
made up
cerebral
of three parts:
(i)
The
the
hemispheres and
inter-brain, comprising the thalami and their associated nuclei, which with the former constituted the fore-brain; and (3) the
mid-brain
(Figs.
17,
18,
and
which
27).
fills
The cerebrum
is
an ovoid
and
rests, below, upon the floor of the cranial cavity in the anterior and middle fossas and upon the tentorium cerebelli over the
Viewed from above, it is sufficiently 2). a sphere; and, being divided in the median line by the longitudinal fissure, the lateral halves are called hemisposterior fossa (Fig.
round
to suggest
the frontal pole, and the most posterior is the occipital pole (Fig. 22). In the floor of the longitudinal fissure of the cerebrum the corpus callosum can be
pheres.
The most
anterior
point
is
seen joining the hemispheres together; and beneath it, concealed from view, are the fornix and anterior commissure. 'Those are
the connecting links, proper, of the hemispheres (Figs. 34, 36 and The latter form Inferior to them is found the inter-brain. 37).
an additional union
of the hemispheres, as
may be
seen by viewing
SO
51
the base of the brain. Just caudal to the inter-brain is the midbrain which occupies the tentorial notch of the dura mater; and, situated in the median line, is so overhung by the cerebral hemis-
its
anterior surface.
It
resembles the
rhombencephalon.
44)-
In studying the gross structures of the cerebrum it is most convenient to divide it into its embryologic divisions, viz., the
fore-brain
SECTION
I.
i.
End-brain
Commissura Hippocampi
Fore -bra in
(Fornix).
Thalami
2.
Inter-brain
Mammillary Bodies
In order to
fix
and relations of the gross structures of the fore-brain it is necessary to study in detail the topography of the exterior surface and the interior surface. It is that with which the present section deals. For the minute anatomy of the cerebral structures, see
Section III of the Cerebrum.
composed
of a thin
sheet of gray matter varying in thickness from one-sixth to onequarter of an inch. That gray matter forms a bark-like cover-
ing for the underlying white substance and is, therefore, called the cortex (Figs. 34 and 35). It is thrown into irregular elongated
52
folds
THE CEREBRUM.
named
convolutions, or gyri,
relative
by deep amount of
is
linear depressions,
cortical substance.
The
increased in bulk to
cerebrum (DeReg-
ibus).
The name fissure is properly applied, first to those deep furrows which represent clefts between embryonic vesicles, viz., the median, verticle cleft between the cerebral hemispheres, and the two arched clefts, one between the cerebellum and the cerebral
hemispheres and the other between the cerebellum and the posterior surface of the medulla oblongata (Figs. 19 and 20); and,
second, the deep linear depressions in the cerebral hemisphere
on the
All other
The
The
basal
surface
comprises
the
orbital
and
tentorial areas,
separated by the stem of the fissura laterah's cerebri (Sylvii). The convex surface is separated from the medial surface by the supero-
superciliaris).
The
medial
border
(margo
orbitalis medialis)
separates the
the medial occipital border (margo occipitalis medialis) divides the medial surface from the tentorial area of the basal surface
(Figs.
19,
22
and
26).
related to
surface of the cerebral hemisphere (fades convexa two very extensive fissures, viz., the longit-
udinal
The longitudinal
is
fissure of the
cerebrum
the vertical
median
FISSURES
53
FISSURES
cleft
5$
and
22).
corpus callosum. The cerebrum is bellum by the transverse fissure of the cerebrum
(fissura
and 6). This forward above the mid-brain, and terminates between the inter-brain and the fornix, where
transfersa cerebri, Figs. 20, 27,
i
fissure continues
in the
it
cerebrum
is
continuous,
by
its lateral
hem-
ispheres.
occupies the posterior part. The anterior part of the transverse fissure contains the chorioid tela of the third ventricle.
The tentorium
There are three great furrows in the convex surface of each cerebral hemisphere which form interlobar boundaries and constitute very important landmarks: The fissura cerebri lateralis, the sulcus centralis, and the sulcus occipito-parietalis (Figs. 22
and
23).
The
It
cerebri
at the
same name
runs outward between the frontal and the temporal lobe, along
;
the lesser wing of the sphenoid bone and, turning upward, on the convex surface, it divides three- fourth inch behind the Sylvian point
into
an anterior horizontal, and anterior ascending and a posterior ramus (Fig. 23). Into the frontal lobe project the small anterior
rami.
They
are separated
by the
Below the
anterior
horizontal
ramus
is
a knuckle of the
same
frontal gyrus
which
and, between the ascending and posterior located the pars opercularis, constituting the connecting
orbitalis;
gyrus
between the anterior and posterior central gyri. The forms the frontal part of the operculum
The operculum (operculum, a cover) (pars frontalis operculi). covers the island. The posterior limb of the lateral cerebral fissure separates the temporal lobe from the parietal. Near
the crotch and within the fissure
is
line
drawn from the Sylvian point, one and one-quarter inches behind the zygomatic process of the frontal bone and one and a half inches above that of the temporal, backward to the subparietal
56
THE CEREBRUM.
lies
point, three-quarters of an inch below the tuber parietale, directly over the posterior ramus of this fissure.
Centralis (Rolandi, Figs. 22, 23, 24 and 27). Beginning just above the posterior limb of the lateral cerebral fissure, is the central sulcus, which extends upward and backward
Its upper extremity about half an inch (or 5.7 per cent.) behind the middle of a line drawn from the nasal eminence to the external occipital
is
The Sulcus
protuberance.
With
this
sagittal
The
average Rolandic
is three (Cunningham). and forms the boundary between the frontal and the parietal lobe. It is developed hi two parts
71
7'
The
sulcus centralis
and
a superior third and an inferior two-thirds, which join at an angle open backward, called the genu superius ; both parts may present
an anterior concavity.
two parts
Often a concealed gyrus separates the genu superius (Fig. 22). This
superior genu is in line with the superior frontal sulcus and marks the probable location of the trunk center and the boundary between
the
less
arm and
There
is
central sulcus;
is
genu inferius, hi the lower part of the in line with the inferior frontal sulcus and
arm area and
the upper limit of the
limit of the
The
Occipito-parietal Sulcus.
(Sulcus occipito-parietalis).
to the
If the line
on the
back
sagittal
meridian
its
The
sulcus
end marks the location of the occipito-parietal sulcus. is located one- sixth of an inch above the lambda in
the adult,
and
is
to
occipital pole.
is
The
situated
it is
on the medial surface of the cerebral hemisphere, hence, divided into an internal part and an external part which are
continuous through the supero-medial border (Figs. 20, 22 and 28). To the extent of its depth, which is about one inch, the external
occipito-parietal sulcus separates the occipital
from the
parietal
Cunningham
FISSURES
57
(U
O.
ll
.5
c c
11
"
.
F*H
~
&.
Ui
J)
o.
.213
-a
iu
>
^23 -
C^
"2
M II -"
59
considers the occipito-parietal sulcus a true fissure because in the embryo it produces a ventricular eminence, though it disappears
during development.*
The
polar region of the hemisphere and forms a part of all three surfaces (Figs. 22, 28 and 26).. On the convex surface, it extends
as far
bounded behind by the stem of the surface, lateral cerebral fissure and the anterior perforated spot; and it is limited posteriorly by the sulcus cinguli on the medial surface
on the basal
is
On
the convex surface, the frontal lobe has the following sulci
(Figs. 23
and gyri
and
24):
Superior precentral (s. prascentralis superior) Inferior precentral (s. praecentralis inferior)
.
j
^
Superior frontal
Inferior frontal
(s.
(s.
(s.
Middle
.
frontal
(s.
frontalis
medius)
Paramedial
paramedialis).
(g. centralis
Anterior central
anterior)
j
I
Middle
The
sulcus
precentral sulci
(Fig.
23)
end of the
sulcus
and the ascending ramus of the lateral fissure of the cereThey form the anterior boundary of the anterior central The superior frontal sulcus and the inferior frontal sulcus gyrus.
brum.
are
respectively continuous with the corresponding precentral sulcus from which they trend downward and forward parallel
*The name
of this sulcus
is
60
THE CEREBRUM.
with the supero-medial border of the hemisphere. They separate from each other three gyri of nearly equal width, viz., the superior, middle and inferior frontal gyri (Fig. 24).
superior frontal gyrus is incompletely divided in the brain by an interrupted sulcus, called the sulcus paramedialis (Fig. 23) which is located near the supero-medial border
of the hemisphere
The human
and is said by Cunningham to be better developed human race and to be rare in the higher
series of
middle frontal sulcus (s. frontalis medius, Fig. 23) partially subdivides the middle frontal gyrus into an upper and a lower part. The middle frontal sulcus, not found below the anthropoid apes
(Cunningham),
furrow,
the
is
best
marked
The
posterior
end,
the
foot, of the
and
It
hemisphere
The
is
human
brain, especially in the left hemisphere of right-handed people. It is deeply cleft along its lower border by the anterior ascending
and and
anterior horizontal
rami of the
cerebrum
is thus divided into a pars orbitalis, situated below the anterior horizontal ramus, a pars triangularis, inclosed between the anterior horizontal and ascending rami, and a pars basilaris, located
lateral fissure
and
the inferior precentral sulcus. The pars basilaris constitutes the foot of the inferior frontal gyrus and is continuous with the gyrus
centralis anterior;
on the
left
side
it
The
anterior portions of the superior middle and inferior frontal gyri comprise a psychic center, center of attention, volition, inhibition, etc.,
The
"of abstract concept" (Mills). anterior central gyrus (g. centralis anterior}
lies
between
the precentral sulci and the central sulcus. It is joined to the posterior central gyrus by the paracentral lobule, above the
LOBE.S
6l
Sjl*
S O
^
tctj
a
_
s|
c_)C/3
g
""*
<^tn^2 O
***
*
.
8-i
Ja
u
_-
g"c.2s "" Vl ?^
rrt
^3
.2 f rt
d a.S'C U <W
O
^
^5
3^^
&
and by the
below
it.
The
fronto-parietal part of the operculum, anterior central gyms, together with the para-
human
cerebrum.
(2) The parietal lobe (lobus parietalis) is situated behind the central sulcus and above the posterior limb of the lateral
fissure of the
cerebrum
(Figs. 22
and
28).
From
the posterior end of the latter to the occipito-parietal sulcus the lobe is separated from the temporal, below, and the occipital,
behind, by an imaginary line. This imaginary line runs backward parallel with the infero-lateral border of the hemisphere to the boundary of the occipital lobe; and then, obliquely upward
toward the supero-medial border in a line drawn from the preoccipital notch to the occipito-parietal sulcus. Extending over
the supero-medial border, the lobe on the medial surface
is
inclosed
between the occipito-parietal sulcus behind and the marginal part of the sulcus cinguli in front, and is bounded antero-inferiorly
by the subparietal
sulcus.
the convex surface of the hemisphere the parietal lobe possesses the following sulci and gyri (Figs. 23 and 24).
Interparietal
(s.
On
post-centralis inferior;
Superior post -central (s. post-centralis superior) Horizontal limb (ramus horizontalis)
Sulci
Upturned ends
of
(f.
Lateral fissure
cerebri lateralis)
(s.
temporalis superior)
(s.
temporalis medius).
Posterior central (g. centralis posterior) Superior parietal lobule (1. parietalis superior)
Inferior parietal lobule
(1.
parietalis inferior)
Gyri
Supramarginal
(g.
supramarginalis)
Angular
(g. angularis)
interparietal sulcus (Figs. 20 and 23) is the only one belonging to the parietal lobe. The inferior and superior post-
The
64
THE CEREBRUM.
central sulci, constituting its anterior parts, are parallel with the
and are located a half or three-quarters of an inch separated from the central sulcus by the gyrus centralis The post-central sulci are often not continuous. The posterior. inferior is about twice the length of the superior, hi this resembling
central sulcus
behind
it,
the central sulcus, and usually it is joined at horizontal limb of the interparietal sulcus.
of the sulcus lies about
its
The
an inch below the supero-medial border of the hemisphere with which it is parallel; it separates the superior parietal lobule from the inferior parietal lobule and is continued as ramus occipitalis into the occipital lobe where it bifurcates.
The
posterior central gyrus reaches from the posterior limb upward and backward, between the central
sulci, to
and post-central
(Fig. 24).
It is
ends of the central sulcus by superficial annectant gyri (gyri transitivi) and sometimes is connected with it by a buried gyrus
(g.
from the
profundus transitivus) which, deeply, separates the superior inferior part of the central sulcus. The annectant
gyrus which closes the central sulcus superiorly and links together the central gyri is the paracentral lobule (lobulus paracentralis) the fronto-parietal part of the operculum joins them below the
;
The posterior central gyrus and paracentral central sulcus. lobule constitute the receptive area of common sensation, the somasthetic area, so far as it extends on the convex surface.
The superior
parietal
supero-medial border of the hemisphere from the superior postcentral to the occipito-parietal sulcus.
inferior parietal lobule
sulcus; posteriorly,
it
by the horizontal part of the interparietal is joined to the occipital lobe by a curved
annectant gyrus, called the arcus occipito-parietalis, which closes the superior end of the occipito-parietal sulcus; and, over the
supero-medial border, it is continuous with the praecuneus of the medial surface. In the praecuneus and the superior parietal
lobule Mills locates the stereognostic center (Figs. 56 and 57). The Inferior Parietal Lobule. The inferior parietal lobule is incompletely divided into two or three gyri. Named from
6s
<
^
h
O 60
vi
t>
>
CB *!8
a1 "
a * c-.2 M'C
fc
.2
ao
67
before backward they are as follows: The supramarginal, the angular and the post-parietal (Figs. 20 and 24).
The supramarginal gyrus arches over and closes the upturned end of the posterior ramus of the lateral fissure of the cerebrum The anterior segment of the arch is continuous with (Fig. 24).
the posterior central gyrus and is comprised in the psychic-sensory area; the posterior segment of the arch fuses with the angular
gyrus,
behind,
memory
center
temporal sulcus (Figs. 20 and 24). The angular arch is in direct continuity with the superior and middle temporal gyri and the posterior segment of it is continuous with the post-parietal gyrus
when
that gyrus
is
present.
post-parietal gyrus is present only when the middle temporal sulcus bends upward and terminates in the parietal lobe; in which case this gyrus curves over and closes that sulcus
(not figured).
It
The
middle temporal gyri and also blends with the superior occipital
gyrus.
The
the
left side,
angular, post-parietal and superior occipital gyri on according to Mills and others, constitute the center
for visual
memories.
The
studies of A.'
W. Campbell
render
it
probable that the visual cortex does not extend into the parietal lobe of man at all. The receptive visual center for macular vision
(vision in the
macula lutea
of the retina)
is
likewise located in
the angular or post-parietal gyrus by Mills, but it is probably situated on the medial surface of the occipital lobe; unlike the memory center, this is present in both hemispheres.
(3)
Occipital Lobe.
(Lobus
occipitalis,
Figs. 20,
22,
24,
26
and
28).
The
hem-
isphere.
tinuous,
With the parietal and temporal lobes it is directly conbeing marked off from them by an imaginary line drawn
preoccipital notch to the occipito- parietal sulcus.
from the
sulcus,
This
on the convex surface, bounds it to the extent of about an inch; rarely, the external perpendicular sulcus bounds it in front. On the medial and basal surfaces of the hemisphere the
68
occipital lobe extends
THE CEREBRUM.
from the occipito-parietal sulcus to the preoccipital notch, and is separated from the temporal lobe on the basal surface by an imaginary line drawn from this notch toward the posterior end of the corpus callosum, to the isthmus of the
limbic lobe.
The
form of a triangular
pyramid whose borders are the supero-medial, the infero-lateral and the medial occipital borders of the cerebral hemisphere. Those borders meet at its apex, the occipital pole.
The
occipital lobe is
pearance at
fissures
somewhat rudimentary in man, though and men (Cunningham). It makes its apthe fourth month in utero, and is distinctly outlined by
three surfaces at the sixth month,
in the adult ape's brain
on
all
when
it
resembles
the
same lobe
(Cunningham Memoirs).
month
from the human brain on the convex and basal surfaces; hence,
artificial boundaries in the adult. Its rudimentary character probably accounts for the great variability in the sulci and gyri of the occipital lobe. On the convex surface they are usually
the
as follows:
(
Ramus
Sulci
<
Transverse occipital
Lateral occipital
(ss.
occipitalis transversus)
occipitales laterales).
|
|
'
Superior occipital (gg. occipitales superiores) Lateral occipital (gg. occipitales laterales).
to the
lateral occipital sulcus is the only one properly belonging convex surface of the occipital lobe (sometimes there are two of them). It divides that surface almost equally into a
r
The
The superior and lateral gyrus, both of w hich may be double. sulcus begins near the supero-medial border. It follows a meridwhich is nearly parallel with the infero-lateral border of the lobe and runs forward, often presenting one interruption, to the occipito-temporal boundary line where is bifurcates and
ian
forms a short perpendicular sulcus, which represents the affenspalte of the ape (Elliot Smith).
is
When
and
24).
The
occipital limb
of
69
not always
from the supero-medial border of the hemisphere, it passes the occipito-parietal sulcus, from which it is separated by the arcus occipilo-parietalis, and bifurcates in the superior occipital gyrus
into
two more or less oblique branches, constituting the transverse occipital sulcus. The medial end of the transverse sulcus may or may not cut the supero-medial border of the lobe. The lateral occipital gyrus (Figs. 20 and 24) lies below the
occipital pole
forward
along the infero-lateral border of the hemisphere to the preoccipital notch. Sometimes it is divided into two lateral gyri by an
inferior lateral sulcus.
It
is
when
by the
The superior occipital gyrus forms the upper half of the convex surface of the lobe (Figs. 20 and 24). It is incompletely separated from the lateral occipital gyrus by the lateral occipital
often divided into two gyri. It is continuous with the post-parietal gyrus around the lateral end of the transverse
sulcus
and
is
it
by
parietalis.
The
latter is
bounds the
occipito-parietal sulcus.
When
upward
at its posterior
continuous
gyri.
As already
to
mentioned,
the
according
Mills,
The temporal lobe (lobus temporalis, Figs. 20, 22, 23, 24, and 26) is that part of the cerebral hemisphere behind the 25 main stem and below the posterior limb of the lateral cerebral
fissure.
It
rests
in
continuous posteriorly
it
the occipital and parietal lobes, from which off only by the imaginary lines already described.
is
marked
medial
On
the basal
its
70
THE CEREBRUM.
is
border
separated from the limbic lobe by the collateral fissure and by a short furrow, which is shallow in the adult human brain, The temporal lobe is attached called the sulcus ecto-rhinalis.
to the
free surfaces
hemisphere posteriorly and medially; but it presents three a superior, an external and an inferior which meet
of the temporal lobe forms the inferior
wall of the lateral cerebral fissure (Fig. 25). It looks somewhat medianward toward the island and constitutes the temporal part
of the operculum.
It is
circular
sulcus
(s.
circularis
separated from the island by the inferior insula). Anteriorly, the superior
surface of the temporal lobe is smooth; posteriorly, it possesses one or two shallow transverse temporal sulci which feebly outline two or three gyri, called the transverse temporal gyri of Heschl
(gyri
temporales
transfer si).
These transverse
gyri
belong to
the receptive auditory center (Figs. 31, 35, 36 and 54). The external surface of the temporal lobe presents the fol-
Superior temporal
(s.
temporalis superior)
J
}
Middle temporal
Inferior temporal
really
(s.
(s.
This
is
on the
Superior temporal
(g.
temporalis superior)
Gyri
<j
Middle temporal
(g.
temporalis medius)
The superior temporal and middle temporal sulci (Fig. 23) divide the external surface into three nearly equal gyri; they run
parallel
The
upward
posterior extremity;
it
terminates in the concavity of the angular gyrus. temporal sulcus is usually an interrupted one. It
The middle
may
present
an upward curve
tion
at its posterior
;
end which
is
may
continue in
original direc-
The
71
the basal surface, runs interruptedly close to the infero-lateral border of the hemisphere and parallel with it (Fig. 26). It separates the inferior temporal gyms from the fusiform gyrus. The superior temporal, the middle temporal and the inferior
temporal gyri are of nearly equal width (Figs. 24 and 26). They fuse with one another and with the fusiform gyrus at the temporal
continuous with the supramarginal and angular gyri, posteriorly: in its third and fourth
pole.
The
is
fifths
and
center (Barker).
The middle temporal gyrus fuses at its posterior end with the angular gyrus and either with the post-parietal or the superior Along the superior temporal sulcus, in the middle occipital. two- fourths of the superior and middle temporal gyri is the center
for auditory memories, the psychic auditory center.
is
This center
handed people (Figs. 54 and 56). The inferior temporal gyrus forms the infero-lateral border It is continuous with the of the hemisphere (Figs. 24 and 26).
in the left hemisphere of right
occipital
lateral
occipital
and post-parietal
gyrus and sometimes, also, with the superior If this gyrus and the lower gyri.
each fourth, according to Mills, belongs to a definite center. From behind forward they are the center of orientation; the center
of equilibration (?); the
naming
The
It is situated in the medial wall of (Figs. 25, 26, 31 and 36). the lateral fissure of the cerebrum, between the frontal, parietal
and temporal
life
it
lobes,
it
gradually covers
is
over.
entirely
whose growth, after the fifth month in utero, At the end of the first year of extrauterine concealed by temporal, parietal, and frontal
The island is thus separated from the parts of the operculum. surface of the cerebral hemisphere by a distance of half general
or three-quarters of an inch. It is triangular in shape. Its apex is directed downward and forward toward the fossa lateralis
cerebri,
and
is
lateral fissure
72
THE CEREBRUM.
seen separating the island from the frontal, parietal and
may be
The
circular sulcus
is
where the
gyrus and with the anterior perforated substance, and is on the same level as the orbital area of the basal surface. The imaginary
line separating the anterior perforated substance
is
is
six gyri
(s.
centralis insulae).
j \
'
Long
(gyrus longus
furcalis).
The
island
and runs obliquely upward and backward dividing the lobe into two lobules (Fig. 25). It is in the same transverse plane
as the central sulcus (of Rolando). In front of it, is the precentral lobule composed of the short gyri and continuous with the frontal lobe; the post-central lobule lies behind it and is in
continuity with the parietal, temporal and limbic lobes. The gyri breves insulae, three or four in number, are separated
shallow, furrows which diverge upward and backward from the smooth apex of the precentral lobule (Fig. 25). They are to the orbital operculum by a short annectant gyrus (gyrus joined
by
connection under the circular sulcus with the foot of the inferior
frontal gyrus suggests a participation in the speech center,
and
which
results
from
Gyrus Longus
It lies
sulcus of the island and trends obliquely backward and upward. Posteriorly, it bifurcates for a short distance forming two short
branches which are continuous under the circular sulcus with the
73
and the limbic lobe, comprising the rhinenand a part of the neopallium, also belong to the cerebral cephalon hemisphere; but no part of either can be seen on the convex surolfactory lobe
The
and
18).
The
inter-brain,
hypothalami and the cerebral hemispheres; and, second, of the which embraces the pars mammillaris hypothalami.
It is completely exposed only when a section is made through the mid-brain and the rhombencephalon removed (Figs. 21 and 26). This should now be done with a thin, moistened brain-knife.
Make
at a right
Now
notice,
first,
the section
structures
median
the
The latter form the very large peripheral zone. The base of the cerebral hemisphere extends from
to the occipital pole.
composed bounded by the medial orbital and superciliary borders and, posteriorly, is made up of the tentorial area, which is bounded, laterally, by the infero-lateral border, and, medially, by the chorioidal fissure and the medial occipital margin of the hemisphere (Fig.
;
In front
it is
26).
The
orbital area
lobe and of the island, and the whole olfactory lobe; while the inferior surface of the temporal and occipital lobes, and the gyrus hippocampi and fascia dentata of the limbic lobe are included in
the tentorial area.
(Fig. 26).
on the
separated from
fellow
by
and
is
bounded behind
74
THE CEREBRUM.
by the lateral fossa and lateral fissure of the cerebrum, overlapped by the temporal lobe. More accurately, the posterior boundary is the anterior perforated substance and the anterior part of the
circular sulcus.
is
The orbital lobe is concave transversely and divided by the triradiate or H-shaped sulcus orbitalis, made up of the medial orbital, the transverse orbital and the lateral orbital
sulci;
inal fissure
and by the olfactory sulcus, which is close to the longitudand parallel with it. Five gyri are thus formed
:
Medial
Gyri
Lateral orbital
not constant
Anterior orbital
Posterior orbital
anterior)
The gyrus
surface.
It is
rectus (Fig. 26) forms the medial border of this separated from the medial orbital gyrus by the
lie
Over
it
gyrus rectus
Broca)
same
It
and the
island.
anterior and posterior orbital gyri lie within the H-shaped orbital sulcus separated from each other by the transverse orbital
sulcus.
The
The former
is
circular sulcus
likewise continuous with the posterior end of the lateral gyrus orbital gyrus and with the orbital portion of the inferior frontal.
The
is
when
the
H-shaped
sulcus.
It
is
The Island
Surface
(Fig. 26).
If the an-
The
circular sulcus
75
Fig. 26.
Base of fore-brain and cut surface of mid-brain, Right temporal pole cut away, to show inferior surface of the island. (Original.}
Sulcus parolfactorius anterior, b. Sulcus parolfactorius posterior, c. Olfactory bulb. Olfactory tract, e. Olfactory striae, f. Area parolfactoria. g. Trigonum olfactorium. Substantia perforata anterior, i. Gyrus subcallosus (peduncle of corpus callosum). k. Optic tract. I. Tuber cinereum. m. Infundibulum. n. Corpus mamj. Optic chiasma. millare. o. Substantia perforata posterior, p. Aqueductus cerebri. q. Quadrigeminal colliculus. r. Corpus pineale. s. Splenium.
a.
d.
h.
77
sides and separates it from the posterior orbital and from the temporal lobe, behind. Laterally it is separated from the frontal and the parietal parts of the operculum by an antero-posterior cleft continuous with the lateral
on two
gyrus, in front;
cerebral fissure.
The
insula
is
of transition
continuous with the anterior perforated substance, from one to the other is called the
Rhinencephalon.
surface.
It
is
The
rudimentary in man.
Many
connected
parts
make
it
up.
It is
torius posterior.
divided into two parts by the sulcus paroljacThese are designated as the pars anterior rhin-
The
pars
anterior of the rhinencephalon embraces, first, the olfactory lobe, which is made up of the olfactory bulb, tract, triangle and the medial and intermediate striae; and second, the area parolfactoria.
Olfactory Lobe.
is
(Lobus olfactorius}.
There
is
studied only on the basal surface of the fore-brain. That is the Belonging to the pars anterior rhinenolfactory lobe (Fig. 26).
cephali,
it
comprises
many
calling
them the
olfactory lobe
connected parts; and the reason for is found in the lower animals
it
and
in the
exists as a
prominent hollow
18).
and
Bulbus olfactorius
Tractus olfactorius
Olfactory
Lobe
Trigonum olfactorium
Stria medialis
Stria-
intermedia.
is
The
olfactory
bulb
(bulbus
olfactorius)
an ovoid mass of
brain matter about half an inch long, one- sixth of an inch wide and a quarter of an inch in vertical diameter (Fig. 26). It is
lodged in the olfactory sulcus of the frontal lobe and rests upon
the cribriform plate of the ethmoid bone through which it receives the twenty or thirty olfactory nerves. The center of the bulb
78
is
THE CEREBRUM.
formed by a gray core derived from the ependymal lining of the embryonic ventricle. The gray core is surrounded by a white
of
sheath
medullated
fibers
running longitudinally;
posterior
form the olfactory tract. Five layers of gray substance thicker on the ventral side, surround the white sheath and constitute the surface of the bulb. The gray subto the bulb these fibers
stance forms the terminal nucleus of the olfactory nerves and gives origin to the fibers of the olfactory tract.
Olfactory Tract (Tractus olfactor ius).
in section, slightly
The
tract
is
triangular
line in
width
concealed in the olfactory sulcus, and (Fig. 26). is a continuation backward of the medullated mitral axones which
It is partially
posterior end the olfactory tract divides into three striae lateral, intermediate and medial, two of which are easily seen. These stria olfactories
At
its
are continuous with the three angles of the tract. The lateral and medial striae diverge and inclose the olfactory triangle between
them.
The
lateral
stria
(stria
lateralis]
courses outward
and
in the
uncus
hippocampal gyrus.
lateral
terminates in the rudimentary gyri, circumambiens and semilunaris, which form the anterior end of the hippocampal
gyrus.
The
lateral
stria
bends sharply perforated space. toward the median line, and runs between the triangle inward,
stria (stria medialis)
The medial
and parolfactory area (of Broca). Its fibers turn into Broca's area and the gyrus cinguli, chiefly, but some of them enter the
triangle,
unite the opposite ends of the gyrus fornicatus. From the dorsal angle of the olfactory tract, a bundle of fibers proceeds into the triangle and frontal lobe, constituting the intermediate
The
Upon
commissure
where
79
down
ward and outward toward the uncinate region of the limbic lobe. The Olfactory Triangle and the Parolfactory Area (of Broca). The triangular portion of the cortex between the medial and
lateral olfactory striae, called the triangle (trigonum olfactorium)
is
stria
continuous medially with the area paroljactoria. The medial marks the boundary between them (Figs. 26 and 27). Both
are
bounded behind by the sulcus par ol) actor ius posterior (transverse part), and the oblique part of the same fissure separates the parolfactory area from the gyms subcallosus (peduncle of the
corpus
in front
callosum).
The
area parolfactoria
(Brocae)
is
limited
by a slight curved depression, the sulcus parolfactorius anterior. On the medial surface, this area joins the gyrus cinguli. The anterior perforated substance (substantia perforata anterior}
of the pars posterior rhinencephali requires further
26).
It
is
mention
(Fig.
fault groove,
in direct con-
The
optic
tract
bounds
it,
postero-medially. Laterally, it forms the limen insults in the floor of the fossa cerebri lateralis, where it is overlapped by the temporal lobe. Superiorly, it is continuous with the base
Coursing along the inner and outer border of the anterior perforated substance are, respectively, the
of the
corpus striatum.
stria,
perforations of this area are for the antero-lateral ganglionic arteries. Tentorial Area of the Basal Surface (Figs. 26, 24 and 28).
The
From
bral
hemisphere presents three nearly parallel gyri, viz., the inferior temporal gyrus, which forms the infero-lateral border;
the fusiform gyrus, the middle one, and the gyrus hippocampi which lies next the mid-brain. The last belongs to the gyrus
fornicatus of the limbic lobe;
it
is
the lingual gyrus, which forms a part of the medial occipital border of the cerebral hemisphere. The fusiform and inferior
8o
occipital lobes.
THE CEREBRUM.
These two lobes are
directly continuous
with
each other on their inferior surfaces, and are only separated arbitrarily by an imaginary line drawn from the preoccipital notch
to the anterior
fissure.
They
separated from the gyrus hippocampi; the ectorhinal sulcus (s. rhinalis) and the anterior part of the collateral fissure lie
arid the
w hile
r
separated from the gyrus cinguli, of the limbic lobe, by the anterior The fissures and sulci of the tentorial area calcarine fissure.
are the following:
Chorioidal fissure
(f.
chorioidea)
(f.
Hippocampal
fissure
(s.
(f.
hippocampi)
Ectorhinal sulcus
Collateral fissure
ectorhinalis)
collateralis)
(s.
temporalis inferior).
The chorioidal
fissure
(/.
chorioidea)
medial boundary of the tentorial area (Figs. 26 and 28). At the surface it appears to be identical with the hippocampal fissure;
upon looking deeper, the two are found to be separated by the fascia dentata and the crus of the fornix. This fissure is separated from the inferior horn of the lateral ventricle, only by a layer of epithelium, 'derived from the roof plate of the telencephabut,
lon.
contains the chorioid plexus of the inferior horn. Hippocampal Fissure. (F. hippocampi, Fig. 28). Along the
It
is
the cres-
known
by
as
in front
closed
The fissure the hippocampal fissure. It extends backward to the the uncus.
it
is
con-
tinuous with the furrow behind and above the corpus callosum, called the callosal sulcus. The hippocampal is a true fissure as
it
indents the whole ventricular wall the long ventricular eminence produced by it is the hippocampus seen in the inferior horn of
;
Ectorhinal Sulcus.
Midway
(Incisura temporalis, Figs. 26 and 29). between the temporal pole and the hook-point of the
is
hippocampal gyrus
81
of the rhinen-
boundary
cephalon in animals with highly developed sense of smell. It man the boundary between the hippocampal and
fusiform gyri.
is
the
Fissura Collateralis
(Figs. 26
and
28).
The
collateral fissure
extends in a somewhat curved course from near the temporal Its anterior two-thirds separates pole almost to the occipital pole.
the hippocampal from the fusiform gyrus; its posterior one- third completes the medial and upper boundary of the fusiform gyrus
from the gyrus lingualis. Inferior Temporal Sulcus (Fig. 26). Only one sulcus belongs within the inferior surface of the temporal and occipital wholly
and separates
it
lobes.
It extends from a point near the occipital pole forward the infero-lateral border of the hemisphere almost to the along
may be
Gyrus Fusiformis.
One
gyrus only
is
found
entirely within
the inferior temporo-occipital region (Figs. 26 and 29). That is the fusiform It extends from near gyrus). (temporo-occipital the occipital pole forward and forms the temporal pole. The posterior nine-tenths of its medial boundary is formed by the
and the anterior one-tenth by an imaginary line and the ectorhinal sulcus; laterally, it is bounded by the inferior
collateral fissure
temporal sulcus.
Gyrus Lingualis.
lateral
The
gyrus lingualis
It is
lies
and
fissure.
hippocampi of the limbic lobe in front. The gyrus lingualis (Fig. 29) forms nearly all of the medial occipital border of the
hemisphere.
It
(Figs. 55 and 57). Limbic Lobe (Lobus Limbus), Inferior Part. The gyrus hippocampi of this lobe is visible on the inferior surface of the fore-brain (Fig. 26). Notice how this crescentic gyrus embraces
6
82
in
THE CEREBRUM.
its concavity the section of the mid-brain. It is separated from the fusiform gyrus by the collateral fissure and the ectorhinal The sulcus; and bounded medially by the hippocampal fissure.
is
flexed
fissure
inward and backward over the and the whole anterior part consti-
uncus hippocampi. The region of the uncus is somewhat irregular and appears to comprise the gyrus circumambiens and the gyrus semilunaris described by Retzius. It represents
-
the greater part of the lobus pyraformis of osmatic mammals and is probably the chief receptive center of smell; it receives the
lateral stria of the olfactory tract
peduncle of the corpus callosum). the hippocampal gyrus be drawn downward somewhat, a rudimentary gyrus may be seen between the hippocampal and
chorioidal fissures.
That
is
It is
continuous
is
with the
and behind
in con-
Like tinuity with the fasciola cinerea and gyrus supracallosus. the hippocampal gyrus it forms a part of the limbic lobe. shall recur to this lobe on the medial surface of the hemisphere. Having studied the basal structures of the cerebral hemispheres,
We
it is
now
in order to
inferior
They occupy
hypophyseal region. They constitute form part of the floor of the third ventricle.
The hypothalamus
mid-brain.
is
the
name
applied
it
Posteriorly
pars optica hypothalami thalami. The former belongs to the telencephalon, the
the diencephalon.
divided into two parts, viz., the and the pars mammillaris hypolatter to
They
Lamina
Pars Optica Hypothalami
cinerea terminalis
Infundibulum and
Hypophysis.
<
Corpora mammillaria.
83
(Fig.
27)
is
most superior
median
structures.
It
is
(cinereum) gray matter closing the end of the neural tube. It extends from the anterior superior surface of the optic chiasma
upward and backward to the anterior commissure, just in front of which it becomes continuous with the lamina rostralis of the
corpus callosum.
in front of
Laterally,
it
is
Behind
is
(Chiasma opticum). The optic chiasma is a quadrilateral sheet of nerve fibers whose anterior angles receive the optic nerves and whose posterior angles give off the optic
Optic Chiasma.
tracts
(Fig.
21).
shape. the lamina cinerea, in the optic groove of the sphenoid bone. The fibers of the optic nerves and tracts compose it. There are
three sets of these fibers, namely, the intercerebral,
The chiasma
With the nerves and tracts attached, it is xis a median structure and is situated beneath
the direct,
and the decussating. A fourth group of retinal and said to be commissural for the
to described,
is
very doubtful.
The
inter-
cerebral fibers are not found in the optic nerves, but form the inferior commissure (Guddeni) which joins together the medial
The
side, respectively
In most vertebrates below mammals, and in the mouse and guinea pig, it is said that the optic fibers all decussate in the chiasma.
Normally
nasal
optici]
in
man and
the higher
mammals,
chiasma direct
(Fig.
67).
The
optic nerves
sclerae of
the front of the chiasma, through the optic foramina; they rise which are connected with the rods and cones
(tractus optici)
The
optic tracts
Each
tract
winds outward and backward around the cerebral peduncle, and divides into a medial and a lateral root (Fig. 43). The roots
84
THE CEREBRUM.
at the
corresponding
geniculate body. the medial root has nothing to do with vision. The fibers of the lateral root (radix lateralis] may be traced to the lateral geniculate body (80 per cent., Von Monokow), to the pulvinar of the thalamus
the 20 per cent, remaining), and the rest to the superior quadrigeminal colliculus. The optic radiation of the capsule conThe medial nects these centers with the medial occipital cortex.
(nearly
all
The
root rises
and ends in the medial geniculate body and thalamus. Its fibers form the commissura inferior (Guddeni). Tuber Cinereum. The posterior border of the optic chiasma Here is continuous with the tuber cinereum (Figs. 21 and 26).
the gray matter
is
thickened
and
centrally
prominent.
The
bulbous infundibulum projects downward from it to rest in the sella Turcica, where it forms the posterior lobe of the hypophysis. The upper end of the infundibulum is hollow (funnel-like). Its
In man the cavity forms the lowest part of the third ventricle. bulb of the infundibulum is solid at maturity, though hollow in
the embryo. It is composed largely of fibrous tissue, notwithstanding the fact that it is developed from the floor of the telen-
cephalon. From the base (superior end) of the infundibulum, the tuber cinereum extends in continuity with the anterior perforated substance on each side of it; and, behind, the corpora
it
perforated substance of the mid-brain. The lamina cinerea and tuber cinereum form the inferior gray
The hypophysis
lobes
closed vesicles, lined with columnar epithelium (in part ciliated), contains a viscid jelly-like material (pituita), which suggested the
old
its
name
The
anterior lobe
is
hollowed out on
posterior surface (kidney-shape) and receives the posterior The hypophysis lobe, the infundibulum, into the concavity.
85
has an internal secretion which appears to have an inhibitive action on growth and has been found diseased in acromegalia.
Corpora Mammillaria
either side of the
(Figs. 21
and
26).
Two
white bodies
median
line,
the pigmented gray matter of the posterior perforated substance. Each is formed superficially by a loop in the columna of the fornix
and is, tfierefore, composed of white substance at the surface. There is gray matter in the interior which forms a medial and a
lateral nucleus (Fig. 46).
of the corpus mamillare also rises the tegmental part of the fascic-
ulus
pedunculo-mammillaris; the basilar portion of the same bundle rises in the lateral nucleus and both parts of this pedunculo-
into
the mid-brain to an
unknown
termination.
is is
Immediately behind the corpora mammillaria perforated substance (Figs. 21 and 26). This
part of the substantia nigra of the mid-brain, perforated for the passage of the postero-median ganglionic arteries. The pons
coursing over
its
behind. Issuing from the inner the large oculomotor nerve; and surface from behind forward, is the smaller
it
is
trochlear nerve.
The
median
must be made through the connecting and the inter-brain, dividing the fore-
Separate the lips of the longitudinal cerebrum; drop the moistened brain- knife down onto the corpus callosum; and make one quick sweep of the
knife toward you. third is produced
Of
by
the surface
now exposed
section.
86
THE CEREBRUM.
It is convenient to study the tentorial area of the basal surface with the medial surface (Fig. 26). In this medial and tentorial
surface there are six important sulci and four fissures (Fig. 28).
Of cingulum
Callosal
(s.
(s.
cinguli)
corporis callosi)
(s.
Subparietal
subparietalis)
(s.
(s.
Occi pi to -parietal
Inferior temporal
occipito-parietalis)
temporalis inferior)
Ectorhinal
(s.
ectorhinalis).
Fissures
Hippocampal
Chonoidal
, .
(f.
,
hippocampi)
.
<
I
.,
,,
.,
(f.
(f.
chonoidea)
collateralis).
Collateral
Sulcus Cinguli (Calloso-marginal Sulcus). Beginning under the middle cut surface and extending in a curve forward, upward,
and backward,
then, turning
just
It
until
it
upward
is the sulcus cinguli (Figs. 27 and 28). the gyrus cinguli and a marginal gyrus, including separates the straight and superior frontal, from one another by its anterior
part; and,
by
its
from the praecuneus. The sulcus cinguli is usually interrupted by one annectant gyrus and often by two. These indicate its
development in three separate parts. At its beginning under the corpus callosum, the sulcus cinguli is almost continuous with a small curved sulcus, which runs
nearly vertically downward, called the anterior parolfactory sulcus (Figs. 28 and 29). Behind that little sulcus there is a
small curved gyrus, the parolfactory area (of Broca), which is continuous with the gyrus cinguli and bounded behind by another
The the posterior parolfactory sulcus. latter separates the area parolfactoria from the gyrus subcallosus. Subparietal Sulcus. About one inch above and behind the
slight
sulcus,
called
posterior end of the corpus callosum there is an irregular sulcus, called the Subparietal, which separates the gyrus cinguli of the
limbic lobe from the praecuneus of the parietal lobe (Fig. 28). The callosal sulcus is the deep furrow between the corpus
"o.
73
.a-^
ITS ? <u
..
SP
l^ia 3
-;3
S
E
|I| ^ m cQE"
.
gj
89
callosum and the gyrus cinguli. It follows the convexity of the corpus callosum and was formerly called the ventricle of it (Fig.
The callosal sulcus, behind the corpus callosum, is continu28). ous with the hippocampal fissure. The occipito-parietal sulcus (Figs. 27, 28 and 20), the internal part, extends downward from the supero-medial border to the middle of the calcarine fissure. The two form a lambda-
shaped
fissure
lambda being
tilted
toward the
anterior
The
the
sulcus.
posterior superior ramus, is the occipito-parietal This latter sulcus cuts the supero-medial border at the
junction of the posterior one-sixth with the anterior five-sixths of that border; it is situated about two inches above the occipital
pole,
and
lies
one-sixth of an inch anterior to the point in the lambda. It separates the parietal lobe from the
cuneus of the occipital lobe. The occipito-parietal sulcus is a deep one. In the embryo the primary occipito-parietal fissure
tricle
produces an eminence in the posterior horn of the lateral ven(Cunningham). It is then a true fissure. But that primentirely disappear,
superficial
fissure.
itive
fissure and the ventricular eminence and the adult sulcus is a secondary and
furrow,
hence
it
is
and not a
the
At the
sulcus
contin-
inferior
gyrus,
gyrus cunei,
fissure,
separates
occipito-parietal
it
is
The calcarine
posterior end of the corpus callosum and runs backward and slightly upward to the low.er end of the occipito-parietal sulcus;
downward
where
it
sulcus occipito-parietalis into an anterior calcarine and a posterior calcarine fissure. These three furrows are continuous with one
human
them from each other: the gyrus cunei the occipito-parietal sulcus from the calcarine fissure separates
QO
THE CEREBRUM.
fissure is separated
by
the
gyrus
cuneo-lingualis
(Cunningham).
The
anterior calcarine fissure indents the medial wall of the posterior horn of the lateral ventricle, producing the calcar avis.
Hippocampal Fissure (Figs. 26 and 28). A crescentic fissure, convex downward begins under the splenium of the corpus callosum
in continuity with the callosal sulcus,
beneath the thalamus to within an inch of the temporal pole, where it is closed by the uncus. It is the hippocampal fissure.
the surface of the temporal lobe this fissure appears to be identical with the temporal extension of the chorioidal fissure; but,
deeply, it lies posterior to that fissure and is separated from it by the fascia dentata. The hippocampal fissure produces a long
ridge in the inferior horn of the lateral ventricle, called the hip(Figs. 28 and 39) describes about two-thirds of a circumference along the concavity of the fornix. It extends from the foramen interventriculare backward over
On
the hip-
pocampal
wall.
fissure,
chorioidal fissure
is
The but separated from it by the dentate fascia. a complete one, involving the whole hemisphere
it
from the
lateral
it, pia mater, plexus of that ventricle. The fissure is peculiar in the fact that between the inter-brain and the fornix there is a transverse slit
ventricle.
The
dipping into
by means
of
which
In
it
is
opposite side.
this
in direct
the border
Collateral Fissure.
26 and 28).
It
temporal pole.
The collateral is a long fissure (Figs. reaches from near the occipital almost to the It is situated below and parallel with the cal-
carine and hippocampal fissures, being separated from the former by the lingual gyrus and from the latter by the hippocampal gyrus.
The
gyrus fusiformis
lies
to this fissure.
Ante-
93
rhinalis]
in osmatic animals.
The
rupted by two annectant gyri and divided into a temporal, an This signifies a persistence occipital and an intermediate part.
of
its
embryonic condition.
The
intermediate
portion,
some-
times assisted by the anterior part, indents the ventricular wall and produces the eminentia collateralis in the inferior horn of
the lateral ventricle.
inferior temporal sulcus is usually a series of indentations rather than a continuous sulcus (Figs. 26 and 28). It is
The
about equal in extent to the collateral fissure from which it is separated by the fusiform gyrus. It is parallel with the inferolateral
border of the cerebral hemisphere. As the inferior temporal gyrus, which forms this border reaches over onto the tentorial area a variable distance, even in the two sides of the same
it
not constant
but
it is
gyri
callo-
(Fig. 28).
The two
from
of
made up
the sulcus cinguli (except its marginal end), the subparietal sulcus, and the anterior part of the calcarine and of the collateral fissures.
Gyrus Fornicatus. The gyrus cinguli and the gyrus hippocampi, joined together at the posterior border of the corpus callosum by the isthmus, and together constituting the gyrus forniThe gyrus cinguli begins anteriorly catus, form the central ring. under the corpus callosum in continuity with the area parolfactoria anterior to the fossa cerebri lateralis;
terminates as uncus just behind that fossa. forms the chief part of the limbic lobe.
The
is
gyrus fornicatus
inclosed between
The gyrus
cinguli
is
the callosal sulcus and the sulcus cinguli, except above the posterior
94
THE CEREBRUM.
it
is
28).
callosum, the anterior end of the gyrus cinguli is continuous with a small vertical gyrus, called the area par ol]actoria (Brocae),
embraced between the anterior and posterior parolfactory sulci and is continuous with the area of the same name on the base of the cerebral hemisphere. This part belongs to
which
is
The posterior end of the callosal gyrus almost separated from the hippocampal gyrus by the anterior
narrow link
left
between
this fissure
and the
splenium of the callosum is the isthmus gyri fornicati. It is claimed by Schafer and others, that the superior part of the gyrus cinguli constitutes a portion of the somaesthetic area; but the
histological investigations of Dr. A.
W. Campbell appear
to dis-
prove such a claim. According to Paul Flechsig, the gyrus cinguli contains the center of taste. He locates the center in the posterior part of the gyrus adjacent to the splenium of the corpus callosum;
it
55).
26 and 28) extends downward and forward, along the hippocampal fissure, from the isthmus to within a half-inch of the temporal pole. Its anterior extremity
(Figs.
separated from the pole by the ectorhinal sulcus, and is bent upward and backward over the end of the hippocampal fissure,
is
gyrus
forming a sharply curved hook, the uncus. The hippocampal is bounded below and laterally by the collateral fissure.
Posteriorly,
it is
The
reflected
uncus hippocampi is continuous with a concealed gyrus, located between the hippocampal and chorioidal fissures, viz., the dentate fascia; a narrow band, which winds over the
part of the
uncus near
its
free point
and
is
Giacomin
the
part of the receptive uncus anterior to the end of the hippocampal fissure, Retzius locates the gyrus circumambiens and gyrus semilunaris, which
area parolfactoria constitute the greater center of smell (Figs. 55 and 57). In the
declares
them
to
contain the end of the lateral olfactory stria and of the gyrus
95
,0
>>C O
O'go'F "
*;"
,\
a;
""73 o c
(D'Cfc
<u
..
p."
"3
53
S'iicj
!&
3-"
.'E
S
.
* | u-|
fdl
<u
Krf'a c
g s a
"H M
als aoii: K
fe
w
cs
O
o-a^.y
Q7
the
and he thinks they form the most important part This region represents the receptive olfactory center.
lobus pyraformis of osmatic animals and, according to Elliot Smith (as quoted by Cunningham) it is the only part of the gyrus hippocampi which properly belongs to the rhinencephalon.
Limbic Lobe.
and
The
enumerated as follows:
g.
parts of the limbic lobe may (i) The gyrus fornicatus (g.
fascia
now be
cinguli
cinerea,
hippocampi*);
(2) the
dentata,
fasciola
supracallosal gyrus (longitudinal striae), and subcallosal gyrus (peduncle of the corpus callosum); (3) one- half of the septum
pellucidum; and
lobe
is
(4)
The
limbic
rudimentary in the
human brain.
The
structures enumer-
ated above under No. 2 and No. 3 are but faint representatives of the strong dentate gyrus seen in animals having no corpus
of the corpus callosum encroaches the dentate gyrus. The limbic lobe partially destroys in part belongs to the rhinencephalon. According to the researches of Elliot Smith the posterior inferior part of the gyrus
callosum.
The development
upon and
hippocampi, that part behind the uncus and below the subiculum, and the whole gyrus cinguli belong to the neopallium and not to
the rhinencephalon.
the cerebral hemisphere is composed of five gyri (Figs. 28 and 26). Beginning under the corpus callosum anterior to the area parolto the frontal pole, then along the border to the occipital pole and, finally, along the supero-medial infero-lateral border to the temporal pole, these gyri are as follows
factoria
marginalis) ending behind as lobulus paracentralis which closes the superior end of the central sulcus; the praecuneus inclosed between the
(g.
frontalis superior
marginal end of the sulcus cinguli and the sulcus occipito-parietthe cuneus, located between the occipito-parietal sulcus and the posterior calcarine fissure the gyrus lingualis, which forms much of the medial occipital border and lies between the calcarine and collateral fissures; and lastly, the gyrus fusiformis (temporo-occipital gyrus}, which lies below the collateral fissure and extends from the temporal pole almost to the occipital pole,
alis;
;
98
THE CEREBRUM.
being separated from the inferior temporal gyrus by an interrupted sulcus near the infero-lateral border of the hemisphere.
The inferior temporal gyrus is chiefly on the convex surface. The paracentral lobule contains the motor center for the opposite
foot, just in front of the central sulcus;
that sulcus
(Figs.
is
55 and
the superior part of the receptive som&sthetic area In the praecuneus is a part of the stereognostic 57).
is
center; the
remainder
lingual gyrus, along the calcarine fissure of each hemisphere constitute the receptive visual center for the corresponding halves of both retinae
and perhaps
The
which have
peripheral ring of gyri seen on this surface belongs to lobes their largest exposure on the convex surface of the
Thus seven lobes belong to the exterior cerebral hemisphere. surface of each hemisphere.
1.
Seven lobes
Frontal
Parietal
.
Temporal
Cerebral Hemisphere
Occipital Island (of Reil)
I.
Neopallium
Limbic
Olfactory, etc.
II.
Rhinencephalon
(archipallium)
2.
Basal ganglion
III.
Corpus striatum.
of
the
fore-brain
ventricles
constitute
the
we
notice
In shape that they occupy a wedge-like space (Figs. 30 and 42). the wedge is rectangular; and it stands, base upward, against the
corpus callosum. Its beveled surfaces look toward the hemThe blade is driven downward as if to split the foreispheres.
brain into lateral halves, the edge resting on the medial structures at the base of the brain. The space is inclosed laterally between
99
the diverging internal capsules, which, within the hemispheres, decussate with the lateral extremities of the corpus callosum. The two structures most necessary to an understanding of the
cavities
and
internal capsule
(Capsula Internd). Looking at the base of the brain, we see two broad bands of nerve fibers, the bases pedunculi, issue from the cerebral hemispheres under cover of
Internal Capsule.
the optic tracts and, converging downward and backward, disappear in the pons (Figs. 21 and 45). Traced in their reverse
direction, the fibers of each basis pedunculi enter the
of the
tional
hemisphere cerebrum and are reinforced by a great number of addifibers from the thalamus. The fibers then radiate toward
This
group
The
opens upward and outward and contains the lentiform nucleus; its solid spout, directed toward the pons and
bell of the funnel
medially, is the basis pedunculi. Antero-inferiorly the fibers in the bell of the funnel diverge to opposite sides of the fissura
cerebri lateralis (Sylvii)
hi
its
continuity,
the hiatus Sylvii; otherwise the funnel is complete. As the internal capsule proceeds into the hemisphere, it impales the corpus
striatum in such
manner as to place the caudate nucleus upon its circumference and to inclose within its walls (to capsulate) the
lentiform nucleus.
The
lentiform nucleus
is
separated externally
and upward
therefore,
and
is
flattened
has,
superior and an
inferior lamina, which, posteriorly, are continuous with each other but, anteriorly, are separated by the hiatus Sylvii.
The
section)
inferior
is
lamina
70).
(or inferior
down
it
and
In front
pass outward beneath the lentiform nucleus and, after winding over the inferior horn of the lateral ventricle, terminate
in the parietal, the insular
and temporal
lobes.
IOO
THE CEREBRUM.
Motor Fibers. The inferior lamina of the internal capsule contains most of the temporo-pontal tract (tractus cerebrocortico-pontalis temporalis] which extends from the temporal
and behind the lentiform nucleus to the nucleus and nuclei of the motor cerebral nerves. The interpons mediate tract (tractus intermedius] rises in the lentiform nucleus
cortex beneath
of the
and runs through the anterior part of the inferior lamina, in its course to the substantia nigra and nucleus pontis in which it
terminates.
The
viz.,
(ansa peduncularis) of the thalamus (common sensory) running chiefly to the somsesthetic cortex; and part of the acustic or temporo-thalamic .radiation (special sensory),
which extends from the medial geniculate body to the auditory In part at cortex in the superior and transverse temporal gyri.
least the ventral stalk rises
thal-
from the substantia nigra and the nuclei of the gracile and cuneate columns probably. The fibers of the ventral stalk
of the
amus
by a
thin sheet of gray substance. The lower lamina (inferior of the thalamus) runs for the most part beneath the peduncle lentiform nucleus into the external capsule and terminates in the
temporal lobe and the island. The upper lamina (ansa lenticularis) enters the lentiform nucleus in two vertical sheets that separate
the three zones of that nucleus.
Its fibers to
a large extent
ter-
minate in the lentiform nucleus, whose axones continue the path through the parietal stalk, in the superior lamina of the capsule,
to the somaesthetic area of the cortex.
inferior
horn and the central part of the inferior lamina joins the superior.
the
(or superior ramus) of the internal It is a contains most of the basis pedunculi (Fig. 30). capsule thick and strong sheet of fibers. Often it is considered as the
The
from the basis pedunculi to the corpus callosum, with which it interfan-like,
101
Fig. 30.
Transverse section of the brain, directed from the pons obliquely upward and forward, showing internal capsule, corpus callosum, ganglia and ventricles of the forebrain.
(Original.)
103
corpus striatum
has a free border anteriorly, though imbedded in the (Fig. 32). Upon its medial surface rest the
farther forward.
thalamus, below and behind, and the caudate nucleus, above and It thus separates the thalamus and caudate
bend seen
in horizontal
section near the middle of the superior lamina, called the genu,
divides
it
occipitalis} which tend slightly and form an angle with each other of about 120 degrees. Motor Fibers of Superior Lamina (Figs. 32, 33 and 70). The fibers of the genu and anterior two-thirds of the occipital
part (pars
part of the internal capsule are very largely motor and constitute the pyramidal tract. They may be traced from the motor area
of the cortex (Figs. 54
and 56) through the internal capsule and the middle three-fifths of the basis pedunculi, and on down into
the spinal cord.
They end
and in the motor cerebral nuclei. same spinal and cerebral nuclei other fibers rise which constitute the motor part of the spinal and cerebral nerves. The of the internal capsule contains a motor tract which pars frontalis extends from the frontal cortex through the inner one-fifth of the basis pedunculi to the nucleus of the pons and motor nuclei of
in the gray matter of the cord
From
these
It
is
Lamina
(Figs. 32,
33 and 70).
In both parts of the superior lamina of the capsule there are common sensory fibers which rise chiefly in the thalamus and end
nary sensations.
and 56). They convey ordiIn the frontal part is the frontal stalk of the thalamus, which ends chiefly in the caudate and lentiform nuclei, though some of its fibers reach the frontal cortex. The tract is
in the somaesthetic cortex (Figs. 54
relayed in the lentiform nucleus, whence it is continued to the somaesthetic cortex by way of the parietal stalk, which is entered
The
superior lamina contains the parietal stalk of the thalamus, which conveys ordinary sensations from the thalamus to the somaesthetic cortex, chiefly to the posterior central gyrus.
1.04
THE CEREBRUM.
of the
common
show
sensory fibers.
that
it
The
up
(alpha and gamma) from the globus pallidus and four from the thalamus. proceed According to Flechsig there are corticifugal fibers intermingled
of six bundles of fibers,
is
built
all
The common
the cortical
fillet.
of the superior
Special Sense Fibers (Figs. 32, 33 and 70). At the junction and inferior laminae of the internal capsule are
The
(radiatio temporo-thalamica] the retro-lentiform part of the internal capsule. The occupy former extends through it from the lateral geniculate body and thalamus to the cuneus and lingual gyrus; the latter rises in the
radiation
temporal
medial geniculate body and ends in the superior and transverse The optic and acustic radiations both contain gyri.
corticifugal fibers
which
rise in
respectively, and run through the capsule to the lateral and medial geniculate bodies and perhaps on to lower nuclei.
which pass through the corpus callosum to the opposite hemisphere other fibers may be traced entire through the same course to the
cortical cells of the opposite side.
bundle of thalamic
fibers
The superior lamina of the internal capsule, proceeding outward and upward into the hemisphere, intermingles with the corpus callosum and enters into the corona radiata. Together with the caudate nucleus, thalamus and stria terminalis (taenia semicircularis),
which
lie
on
its
medial surface,
it
lateral
boundary
thus
Corpus Callosum
(Figs.
34 and 42). The entire roof representing the base of the wedge, is
29, 30,
formed by the corpus callosum. A part of the anterior boundary is also formed by it. The corpus callosum is a thick sheet of
105
107
and a
missure,
being composed chiefly of those medullated cortical axones which end in arborizations about cortical cells of the
opposite hemisphere. It contains some fibers which belong to the internal capsule; and, also, collaterals from capsular and
association fibers.
The
corpus callosum
is
Separating seen in the bottom of the longitudinal about an inch in transverse length.
it is
is
side,
and divided
30 and 34).
median
line
by a longitudinal raphe
(Figs.
Transverse striae are plainly visible. Two longitudinal striae are also found running on either side of the raphe; one next the raphe, the medial longitudinal stria; and the other near the lateral
end of the callosum, the lateral longitudinal stria. The medial and lateral longitudinal striae are imbedded in a thin sheet of gray
substance spread over the corpus callosum; altogether they constitute the
gyrus supracallosus.
border of the callosum, this continuous with the fasciola cinerea and, through that, with the fascia dentata. The gyrus supracallosus becomes the gyrus subcallosus (peduncle of corpus callosum) after it winds around the anterior border of the corpus callosum. As such it is continued
downward between
factory sulcus to the base of the cerebrum, anterior perforated substance to the uncus.
the lamina terminalis and the posterior paroland then across the
downward
hence,
it
is
ward.
Its inferior
riorly
and near
surface (Figs. 29 and 30) is concave antero-posteits posterior border is fused with the body of the
fornix.
the
Anterior to that fusion, it is joined to the fomix, along median line, by the septum pellucidum.
The posterior border (Fig. 29) is flexed downward from the horizontal about forty-five degress. Giving passage to the fibers which join the middle and posterior parts of the hemispheres,
108
THE CEREBRUM.
the posterior border is the thickest part of the corpus callosum. It is on that account called the pad, or splenium.
is
sweeping through 180 degrees of flexion (Fig. 29). It tapers down to a sharp edge, called the rostrum. A very thin sheet-like
extension of the rostrum, called the lamina rostralis, proceeds
backward from the beak and becomes continuous with the lamina terminalis at the base of the brain. Running downward on either side is a low ridge, continuous with the striae longitudinales, which constitutes the gyrus subcallosus. Each gyrus subcallosus,
after passing across the anterior perforated substance, ends in the uncus of the hippocampal gyrus. The transverse fibers of
the rostrum, in the hemisphere, form the floor of the anterior horn
of the lateral ventricle.
(Fig. 29).
is
The down-turned
It joins
the genu.
main body, the truncus. The genu forms part of the anterior boundary of the cerebral cavity; the truncus forms the roof.
Fibers uniting the frontal lobes of the cerebrum pass through the genu, and in the hemisphere, bound the anterior horn of the
lateral ventricle
in front.
Those
fibers
arching for-
of the anterior
forceps minor.
The
forceps major,
composed
from the
splenium which bend backward into the occipital lobe, lies in the roof and inner wall of the posterior horn and produces the eminence
called the bulb (Fig. 38).
overhung by
stria.
is
the gyrus cinguli, which covers the lateral longitudinal Inclosed between the gyrus cinguli and corpus callosum
callosal fissure (ventricle of the callosum).
the
The
lateral extremity
mingles with the superior lamina of the internal capsule and thus stretches entirely across the fore-brain cavities (Figs. 30
and
42).
The boundaries
may be
IOQ
Corpus callosum.
Floor (edge of wedge)
Tegmenta
of mid-brain,
Tuber cinereum,
Infundibulum, Optic chiasma.
Lateral wall (beveled surface) Internal capsule (superior lamina),
Caudate nucleus,
Stria terminalis,
Thalamus.
Anterior wall (border of wedge)
Lamina
terminalis,
Anterior commissure,
Genu
of corpus callosum.
Posterior wall Posterior commissure with cerebral aqueduct beneath Corpora quadrigemina of mid-brain,
it,
The
fore-brain
cavity
thus bounded
42).
is
subdivided by two
35 and
The body
with the chorioid tela of the third ventricle and the roof epithelium of the third ventricle, forms a horizontal partition which divides
the cavity into an upper and lower chamber. The superior chamber is divided into two lateral chambers, the lateral ventricles,
by a double vertical partition, the septum pellucidum. rior chamber is the third ventricle
The
infe-
of the fornix (corpus fornicis, Figs. 29 and 36) is a triangular sheet of fibers, whose base is attached to the under surface of the splenium of the corpus callosum, and whose bifid
The body
to
the
110
the
THE CEREBRUM.
thalami, the third ventricle,
is
together with the chorioid tela and a layer of epithelium. The body of the fornix is produced by the approximation of two
bundles of white
fibers, one belonging to each hemisphere. bundles are the crura of the fornix.
These
The cms fornicis (Figs. 29, 38 and 39) may be traced from the uncus and the hippocampus, its chief origin, upward through the inferior horn and into the floor of the body of the lateral ventricle,
the
where it unites with its fellow of the opposite side in forming body of the fornix. At the apex of the body of the fornix, which is the anterior end, the bundles again separate and become the columnae of the fornix. The crura are united at the back part of the body of the fornix by a few transverse and obh'que fibers which form the lyre, or commissura hippocampi (Fig. 36). The commissure is best seen when the corpus callosum and fornix are viewed from below; its fibers connect each crus of the fornix with the hippocampus and uncus of the opposite side. The columnae fornicis (Figs. 29 and 40), one on either side
pass
down
ventricularia (Monroi)
laria, at
the base of
bounding the foramina interand then descend to the corpora mammilthe brain. On the way down, each columna
behind the anterior commissure, beyond inner part of the thalamus of the
(pars
libera)
passes
which (pars
side. The fibers of the columna fornicis for the most part terminate in the medial nucleus of the corpus mammillare, from which other fibers take their origin, forming the fasciculus mam-
same
millaris
princeps.
is
This
bundle
divides
Y-like;
the
anterior
branch
bundle
(Vicq d'Azyri) and ascends to the anterior nucleus of the thalamus the posterior
fasciculus
;
the
thalamo-mammillaris
the tegmental part of the fasciculus pedunculo-mammillaris and probably ends in the stratum griseum centrale of
is
the mid-brain.
columna of the fornix and, bending runs as medullary stria along the thalamus to the backward, nucleus of the habenula; some of the fibers decussate through
bundle of
fibers leaves the
Ill
the stalk of the pineal body to the opposite nucleus habenulae and constitute the commissura habenularum. The columna
of the
fomix
is
joined by a small fasciculus from the medial root which runs backward to the hippocampus
of the fornix
and uncus.
The upper
surface of the
body
is
convex from
before backward (Figs. 29 and 36). It forms the postero-medial part of the floor of the lateral ventricle. Along the median line,
it is
walled median partition, divides fore-brain cavity into lateral halves, the lateral ventricles.
by the septum pellucidum. 29, 35, 40 and 73), a doublethe superior chamber of the
The
septum pellucidum
fits
is
crescentic in outline.
Its
convex border
corpus
callosum.
Its
concave border
of
commissure, the septum pellucidum is continuous \\ith the gyms subcallosus with which it is associated in development and function.
The septum pellucidum, like the anterior commissure, corpus callosum and fornix, is developed from the thickened upper border of the lamina terminalis and the adjacent medial wall of the
cerebral
These several structures extend upward and backward with the development and rotation of the hemispheres and, together,
roof over the inter- brain.
appears in the
septum and
commonly
The
fore-brain cavity thus embraces four ventricles, viz.: Two lateral ventricles (the ventricles of the hemispheres),
Third ventricle
Fifth ventricle (the ventricle of the septum), and (ventricle of the inter-brain).
Septi Pellucidi.)
ventricle is
the ventricle of the septum (Figs. 35 and 73). The a very narrow, antero-posterior cleft between
it
coincides in
112
extent.
It is situated
THE CEREBRUM.
within the concavity of the corpus callothe lateral ventricles, above and anterior to the
sum between
third ventricle.
It is
Below and
posteriorly
it is
bounded by the
fornix.
not a part of the embryonic brain cavity, but a mere lymph Therefore it does not communicate with any other space.
ventricle,
neural
each of the others being a part of the cavity of the tube from which both brain and cord are developed.
Instead of ependyma, which lines other ventricles, the lining of the fifth is endothelium. A lymph-like fluid fills it.
Ventriculus
La teralis
The hemispheres
30, 35, 41, 42
line,
and
73).
contain the largest of the six ventricles (Figs. Situated one on either side of the median
the ventricles of the hemispheres are very naturally called Each represents a branch of the cavity of the embryonic neural tube (Figs. 17 and 41). In consequence, the lateral ventricles.
all others except the the interventricular foramen (of Monro), each directly By communicates with the third ventricle and through that, indirectly,
fifth.
;
and
sixth.
The foramen
interventriculare
is
of
the fornix (Fig. 29). It extends between the anterior extremity of the third ventricle (the aula) and the junction of the anterior horn with the central part of the lateral ventricle. The lateral
ventricles are lined with
ependyma, which
is
a transparent
mem-
brane composed of two layers when complete, viz., neuroglia and a covering of columnar' ciliated epithelial cells. Over the
thalamus (the part seen in the
plexus, the neurogliar layer
is
lateral ventricle)
absent.
The
lateral ventricle
may be
central part (or body); the anterior horn; the inferior horn;
and
The
is
the ventricle of the parietal lobe of the cerebrum. are its boundaries
:
The
following
Roof
Corpus callosum.
115
chorioid plexus,
Medial wall
External wall
Septum pellucidum.
Internal capsule.
roof inclines upward and outward from the septum pellucidum, the inner wall of the ventricle, to
JL
Fig. 33. Diagram of internal capsule in colors. (Original.) RED, motor; BLUE, common sensory; PURPLE, special sensory.
the superior lamina of the internal capsule, which forms its outer wall. The floor of the central part of the ventricle is formed by
the five parts, as named above, which will in the order given.
now be
considered
is
The striated body (Figs. 30, 31, 32 and 40). the basal ganglion of the hemisphere. It is an ovoid mass of gray matter imbedded, for the most part, hi the cerebral medulla
Corpus Striatum
;
but
continuous below with the anterior perforated substance, and extends above to the lateral ventricle. It measures two
it is
and a
half inches
Il6
transversely, and,
It
is
THE CEREBRUM.
from above downward, one inch and a
half.
placed anterior and external to the thalamus and forms the third of the great divisions of the cerebral hemisphere, viz.,
is
the neopallium, the rhinencephalon and the corpus striatum. It a reddish-gray body, and its streaked appearance is due to the
white capsular fibers which pierce it. The striated body is an important relay hi the motor conduction path and one of less
importance in the sensory path. The internal capsule divides it into two nuclei, namely, the lentiform nucleus (extraventricular part), and the caudate nucleus, which is seen in the boundary of
Anterior to the free borders of the superior and inferior capsular laminae, the two nuclei are united with
the lateral ventricle.
each other, with the anterior perforated substance and with the lower end of the claustrum.
(nucleus
lentijormis]
occupies the
it
cone-like cavity of the internal capsule, separated from the ventricle (Fig. 32).
aft
by whose laminae
It is shorter fore
is
and
It
when viewed
section
in horizontal
31).
In transverse
vertical
is
triangular in shape.
The
formed, respectively, by the superior and inferior laminae of the internal capsule. The external capsule forms the perpendicular
and separates the lentiform nucleus from the claustrum. The a thin sheet of isolated gray matter, found just medial to the island (of Reil). In extent and position, fore and aft, the
latter is
island
is
coincide.
The
lentiform nucleus
subdivided by two white laminae, parallel with its external surThe outer zone, called the face, into three zones. (Fig. 30).
putamen, is deeply pigmented, and, like the caudate nucleus, is of a reddish-gray color; but the two inner zones, having less pigment, are of a pale yellowish tint. They form the globus pallidus.
is
a pear-shaped
inter-
It is the intraventricular capsule (Figs. 30, 32 and 40). part of the striated body and forms a strip of the ventricular floor along the outer wall. The head (capuf) of the caudate nucleus
117
Fig. 34.
Dorsal surface of corpus callosum, cerebral hemisphere cut away to expose it. (Original.)
IIQ
directed forward.
It is
ventricle.
From
downward
and ends
in a
considerable mass of gray substance called the nucleus amygdala, which appears to be an ingrowth of cortex from the region of the
uncus hippocampi (Fig. 32). The caudate nucleus is covered on its ventricular surface by ependyma. The opposite surface, resting against the fibers of the internal capsule, is irregular and
serrated.
stria terminalis (taenia semicircularis, Figs. 30 and 36) It is a band of white lies just medial to the nucleus caudatus.
fibers traversing the floor of the central part of the ventricle
The
and
the roof of
its inferior
horn, but covered by the terminal vein It may be said to rise from the nucleus
amygdalae. Ascending to the ventricle, it passes forward between the caudate nucleus and the thalamus to the interventricular
foramen, where
it
One
of
them accom-
panies the columna of the fornix for a little way. The other, passing over the anterior commissure, descends in front of it and,
;
according to Kolliker, both bundles end in the anterior perforated substance (Cunningham). The thin medial border of the substance covering the vena terminalis, which is attached to the thalamus, is called the lamina affixa; it is present only in the body The lamina affixa is joined by a single layer of of the ventricle. epithelium, the lamina chorioidea epithdialis, to the lateral border
of the fornix.
lateral ventricle.
This lamina invests the chorioid plexus of the If the chorioid epithelium be torn away, the
stria terminalis is called the
t&nia
Thalamus
ventricle,
It
36 and 40).
is
between the
stria terminalis
extends throughout the central part of the ventricle from the A transparent interventricular foramen to the inferior horn.
layer of epithelium, the lamina chorioidea epithelialis, extending to the stria terminalis and representing the hemi-
120
sphere wall, covers
the third ventricle
it.
THE CEREBRUM.
The thalamus
inter-brain.
will
be described with
and
(plexus chorioideus, Figs. 35, 36 and 37) the vascular border of the chorioid tela
It
projects, laterally,
fornix
crus through the chorioidal fissure into the floor of the central part of the ventricle and the inner wall of the inferior
its
and
The epithelium, above mentioned, invests it; and it borders It is called chorioid plexus (chorion, the fornix like a ruffle. a membrane) because it is membrane-like. At the junction of
horn.
the central part and inferior horn of the lateral ventricle the chorioid plexus presents a large skein-like mass called the glomus
The
from the
branch of
and the
postero-lateral chorioidal, a
the posterior cerebral, supply the plexus. The former pierces the temporal lobe and enters the apex of the inferior horn of the
through the transverse and chorioidal cerebrum, following the chorioid tela. The chorioidal vein carries the blood away. At the foramen interventricuventricle; the latter passes in
fissures of the
lare, it is
joined by the terminal vein of the striated body and the veins of the septum pellucidum and forms the internal cerebral
The internal cerebral vein courses backward in the chorioid and unites with its fellow of the opposite side, proximal to which union it receives the basilar vein and then the great cerebral
vein.
tela
;
vein (of Galen), uniting with the inferior sagittal sinus, forms the
straight sinus.
The
is
com-
pleted by the superior surface of the fornix. The horns of the lateral ventricle are three in number; the
anterior, inferior
and posterior
(Figs. 39,
40 and 41).
(cornu anterius, Figs. 35, 36 and 73) projects from the central part of the ventricle forward and outward around the head of the caudate nucleus. It is the ventricle of the frontal
lobe and
is
Its
Roof
Floor
Anterior wall
Genu.
121
Fig-
Horizontal section of cerebrum, cutting splenium and genu of corpus 35callosum, showing lateral ventricles, septum pellucidum, fornix and transverse temporal gyri. (Original.}
123
Septum pellucidum.
Caudate nucleus.
Outer wall
The
posterior
is
horn
and 41)
directed
inward, from the ventricular center into the occipital lobe; and,
like the occipital lobe,
first
makes
its
month
of
embryonic
life.
Its extremity
the posterior calcarine fissure, with which the horn is parallel. The anterior calcarine fissure produces the ridge along the inner wall called the calcar avis. The posterior horn is roofed over
by
fibers
which turn
down
outside the horn and also form part of the external boundary. In the external wall and in the roof and floor is also the optic
well-marked bundle of fibers from the splenium, found passing along the medial border of the roof forceps major, into the occipital lobe. It produces an eminence above the calcar
radiation.
is
anterior extremity of the posterior horn with the beginning of the inferior horn. At the junction of the two is a triangular area, the trigonum collateral. The inferior horn (cornu inferius, Figs. 32, 38, 39 and 41)
The
is
as
the ventricle of the temporal lobe. Its course it follows the of the internal capsule. perimeter
is
crescentic,
It first
runs
outward and backward from the body of the ventricle, then it turns downward, and finally it proceeds horizontally forward and inward to within an inch of the pole of the temporal lobe.
In horizontal section just below the general cavity of the ventricle, the inferior horn is triangular. In that position it has a posterior wall (or floor in the horizontal part), a medial wall, and a curved
antero-lateral wall
which
is
continuous above with the outer wall and floor of the central
part of the ventricle.
The parts found in the walls of the inferior horn ated as follows:
Roof
(or antero-lateral wall)
may be enumer-
Inferior
tail of
caudate nucleus, stria terminalis and amygdala. Floor (or posterior wall) Tapetum
124
THE CEREBRUM.
Eminentia
collateralis
(trigonum collaterale),
Dentate
fascia.
The
ciently described.
They
when
it is
recalled
floor of the central part of the ventricle; the tapetum, the internal
the
cauda,
stria
Beginning at the trigonum collaterale (Figs. 38 and 39) and extending along the outer border of the floor to the end of the
inferior
fissure.
collateral
It is the eminentia collateralis, and is present only when the anterior part of the collateral fissure, as well as the middle The short eminence at the entrance part, is a complete fissure.
trigonum collaterale, is constant presence; produced by the middle division of the collateral fissure. In front of this eminence and internal to it is a
to the inferior horn, called the
its
it is
in
prominent ridge, the hippocampus, which enlarges downward to a lobulated extremity, called the digitations (digitationes hippocampi, Fig. 39). The ridge is due to the infolding of the floor over the hippocampal fissure on the medial surface of the cere-
brum.
The
hippocampus
is
formed
by a lamina of white matter, the alveus, but the deeper part is cortical matter composed almost entirely of pyramidal cell-bodies.
hippocampi) rests in the concavity of the hippocampus, where most of its fibers originate, though a small bundle of them passes beyond it to its origin in the uncus.
o] the fornix (fimbria
The cms
chorioid epithelium (lamina chorioidea epithelialis) (Figs. 62 and 35), -representing the hemisphere wall, forms the 118, floor of the chorioidal fissure and the whole medial wall of the
The
125
Fig. 36.
Horizontal section of cerebrum just below splenium of corpus callosum, snowing commissura hippocampi, fornix, septum pellucidum, the island and
lateral ventricles.
(Original.)
S. C. P. Sul. circu]aris posterior.
127
covers the cushion-like projection (the pulviwhich forms a small part of both roof and
Behind, it is attached to the cms of the fornix, from extends forward to the stria terminalis. The epithelium covers the chorioidal -fissure except at the lower part, where there
inner wall.
which
it
is
a small cleft which forms a communication between the horn and the anterior subarachnoid space. Through the chorioidal fissure a fold of pia mater projects toward the ventricle, and
it
Bounding this part of the chorioidal below and behind, there is a serrated free border of cortex called the dentate fascia. The dentate fascia (Figs. 38, 42 and 62)
hippocampal fissure and with the hippocampus, the gyrus hippocampi and the eminentia collateralis forms an S-shaped fold of the cortex. The S-shape is perfect
folds medially in front of the
when
the anterior segment of a coronal The top of the letter is the dentate
hippocampal
campi;
the
fissure;
open toward the ventricle, is the groove between the hippocampus and the eminentia collower
lateralis.
and Diencephalon.}
The inter-brain
27, 28, 29,
30 and
(diencephalon) is median in position (Figs. It is situated beneath the fornix and the 40).
layer of epithelium extending from the border of the fornix to the stria terminalis. The chorioid tela of the third ventricle only
Laterally, it is bounded by the superior laminae of the internal capsules. The ventricle of the inter-brain is the third in number. The third ventricle, therefore, is located
in the
median plane; and is at a lower level than the ventricles of the hemispheres. Through the interventricular foramina, its anterior part (the aula) communicates with each lateral ventricle, and the cerebral aqueduct connects it, behind, with the fourth ventricle (Figs. 17 and 41). The third ventricle is fissure-like.
128
THE CEREBRUM.
It is a narrow, vertical cleft about an inch in length from before backward and a quarter of an inch broad at its widest part It the thalami, and extends almost to the inferior surface separates of the cerebrum. The roof (Figs. 29, 36, 37, 40 and 42) follows the curve of the fornix and arches from the posterior commissure forward to the anterior commissure. There is a little recess above the anterior commissure and between the columnae of the fornix, bounded in front by the inferior angle of the septum pellucidum, called the recessus triangularis, in which the roof and anterior wall meet. The anterior wall extends from the triangular recess down to the optic recess, at the angle between the lamina cinerea and the optic chiasma. This angle is so named because, on either side of it, there is a lateral extension of the third ventricle between the lamina terminalis and the columna
of the fornix,
vesicle.
which
is
The
the ventricle.
The
first
and
short, stretches
between the optic recess and the infundibulum, in which the floor reaches its lowest point. The distance from the infundib-
ulum
by
is
the
second arch.
but a sixteenth of
aqueduct is spanned and flat. Its posterior extremity long an inch below the posterior commissure; the
It is
The anterior orifice of the cerebral aqueduct separates them. behind to the size of the cerebral
aqueduct, with which it is continuous. The lateral walls (Figs. 28 and 29) are close together throughout. At one point near the
inter-
is
Antero-superiorly, the lateral wall perforated by the interventricular foramen (of Monro). That foramen constitutes the slight separation between the front of the
opens into the lateral ventricle at the junction of the anterior horn with the central part. The ependyma which lines the third ventricle is continuous through
It
the interventricular foramen with the lining of the lateral ventricle. But one layer of the ependyma is present in the roof of the ventricle; that is
The
true ventricles,
occupied by cerebro-spinal
129
Fig- 37-
and
and commissura habenularum, Roof epithelium and pineal body, Chorioid tela and plexuses, Fornix and commissura hippocampi.
Posterior commissure
Fig. 38.
Transverse section of
(Original.)
left
cerebral hemisphere cutting the splenium and floor of the inferior horn of the lateral
c. a. Crus fornicis. b. Fis. hippocampi, Hippocampus, d. Fascia dentata. e. Eminentia collateralis. f. Fis. collateralis. h. Calcar avis. i. Bulb caused by forceps major, j. Tapeturn, k. Radiatio occipito-thalamica. 1. Fasciculus longitudinalis inferior.
Anterior wall
Epithelium, covering
Columnae of
fornix, anterior
commissure, and
Lamina Floor-
terminalis.
Optic commissure,
132
THE CEREBRUM.
-Posterior perforated substance (of mid-brain),
Tegmenta
Posteriorly Ventricle
is
(of mid-brain).
Lateral walls
Thalamus, and
hypothalamic substance,
the back part of the third ventricle and supports the posterior end of the roof epithelium. That band is the posterior commissure (commissure,
posterior,
Columna of the fornix, and Foramen interventriculare between them. Roof. A band of white fibers passes across
Figs. 28
and
40).
It crosses
immediately in front of
the corpora quadrigemina. Beneath it is the anterior orifice of the cerebral aqueduct. The pineal body is above and behind it,
and the commissure fuses with the ventral pineal lamina. The posterior commissure stretches from the central gray substance of the mid-brain on one side, over the aqueduct, to the gray substance of the opposite side and also contains decussating fibers of the medial longitudinal bundle (Heald). The commissure
is
in
third ventricle
and
laterally, is
It is
the superficial layer of the ependyma; but it is, here, the only adult representative of the roof of the diencephalon. The roof epithelium presents two longitudinal folds suspended
in the ventricle.
amus.
The lower
epithelium superiorly; and, dipping down into the longitudinal folds, that inferior layer forms the chorioid plexuses of the third ventricle, which are continuous with those
of the lateral ventricles through the interventricular foramina. At the back part in the middle line, there is a pouch-like evagina-
which develops
pineal recess, in the adult condition. A second evagination occurs just above the pineal recess, which forms the epipineal recess.
Pineal B'ody.
It is
135
cone-shaped body, a quarter of an inch high and one- sixth of an inch in diameter, joined to the roof of the third ventricle by a It is also called the epiphysis. The flattened stalk or peduncle.
pineal body is situated in the floor of the transverse fissure of the cerebrum, directly below the splenium of the corpus callosum
and rests between the superior colliculi of the quadrigeminal bodies on the posterior surface of the mid-brain. It is closely The pineal stalk splits into a dorsal and invested by pia mater. a ventral lamina, which are separated by the pineal recess. The
ventral lamina fuses with the posterior commissure; but the dorsal stretches forward over the commissure in continuity with
The border of the dorsal lamina is thickthe roof epithelium. ened along the line of attachment to the thalamus and forms the
stria medullaris
thalami (pineal
stria).
The
thickening
is
due
to the presence of a
bundle of
fibers derived
of the fornix
Between
the medullary striae, at the posterior end, there is a transverse band, called the habenula. The habenula contains the commis-
sura habenularum, through which the fibers of the striae partially decussate to the nucleus habenulae in the thalamus.
The
body
is
made up
of closed follicles
surrounded by ingrowths of connective tissue. The follicles are filled with epithelial cells mixed with calcareous matter, the brain-sand (acervulus cerebri). Calcareous deposits are found
also
on the pineal
function of
and along the chorioid plexuses. the pineal body is unknown. It is supposed to
stalk
The
repre-
sent a cyclopian eye. In the Hatteria, a New Zealand lizard, it the parietal foramen and presents an imperprojects through fect lens and retina and, in its long stalk, nerve fibers.
tela of the third ventricle (velum interpositum, Figs. 37, 40 and 42) is the double triangular fold of pia mater spread over the dorsum of the inter-brain. It lies underneath
The chorioid
the fornix and the chorioid epithelial lamina which stretches from the body of the fornix lateralward to the stria terminalis.
Its
apex
is
just
its
base,
directed backward,
continuous,
136
THE CEREBRUM.
and cerebellum.
the floor of
its
continuous with the pia on the posterior surface of the mid-brain Each border constitutes the chorioid plexus
and
is
central part.
plexuses of the third ventricle, tion. Between the inferior and superior laminae is enclosed some connective tissue through which the internal cerebral veins run
This layer forms the two chorioid which depend from its median por-
backward
Anterior Wall.
of white fibers
40 and 73) is a very distinct round bundle about an eighth of an inch in diameter. It is seen
epithelium
there
epithelium.
The
bends
down between
the
commissure.
The columnse
of the fornix
bound
the recessus triangularis, in which the roof and anterior wall of the third ventricle meet. The anterior commissure rests
between the
corpus
rostralis of the
it is
callosum, in front.
With the
two structures
developed
the most important connecting link between the hemispheres in vertebrates without a corpus
callosum
(all
below mammals).
hemisphere
its fibers
in the cerebral
the' anterior
It concortex, entering the external capsule. tains two groups of fibers: (i) The anterior group, which is the commissure of the rhinencephalon, called the pars olfactoria;
some
of
and
group, the pars occipito-temporalis. The pars olfactoria probably contains two fasciculi: (a) A commissural bundle which bends sharply downward to the olfactory
(2) the posterior
tracts
and
joins the
(b)
137
Fig. 40.
Horizontal section of cerebrum through genu and below splenium of corpus callosum. Fornix and chorioid tela turned back, to show inter-brain
third ventricle.
of
and
a.
(Original.)
caudate nucleus, b. Stria medullaris thalami (or pineal stria.) c. Chorioid groove, d. Trigonum habenulae. e. Pineal body. f. Tail of caudate nucleus, g. Tapetum. h. Occipito-thalamic radiation, i. Inferior longitudinal fasciculus, j. Anterior horn of lateral ventricle, k. Columna of fornix. 1. Recessus triangularis. m. Anterior commissure, n. Massa intermedia (or middle commissure), o. Posterior commissure, p. Superior quad rigeminal colliculus. q. Posterior horn of lateral ventricle.
Head
139
and amygdala. The pars occipito-temporalis connects the tentorial areas of the two hemispheres together, regions which are not connected by the corpus callosum. In man it is larger than the pars olfactoria. A thin transverse sheet of gray matter,
called the
lamina terminalis,
extends
to the optic
Between the anterior wall of the ventricle (Figs. 17, 28 and 73). the chiasma and the lamina terminalis is a sharp angle which terminates on either side in a small pit, called the optic recess.
is very narrow (Figs. 21 and formed by the interpeduncular structures plus the 27). tegmenta, namely: optic chiasma, tuber cinereum and infundibulum, corpora mammillaria, posterior perforated substance and
The
It
the tegmenta. The last two are portions of the mid-brain; the others belong to the fore-brain with the surface of which we have
The
extend laterally beneath the thalami. third ventricle has its lateral wall formed chiefly by the
all
(Figs. 27
and
28).
Below
aqueduct and called the sulcus hypothalamicus, covered by upturned hypothalamic gray matter
and by the upper part of the central gray substance of the midbrain. The thalamus forms the immediate lateral wall above
this
hypothalamic groove.
its
from
fellow,
The columna of the fornix, diverging downward and backward to the corpus proceeds
mammillare through the medial part of the thalamus. In the ventricle, the pars libera of the columna fornicis is covered by the ependymal epithelium. It bounds the interventricular fora-
men
in front.
Thalamus.
(Thalamus
the great ganglion of the inter-brain. The tant sensory relay station. In it or in the hypothalamic nuclei almost every impulse of general sensation, in its journey to the
The third transferred to a higher neurone. the thalami from each other, except at the separates mid-point where they are joined by the massa intermedia. The
cerebral cortex,
ventricle
is
thalamus
is
situated behind
and medial
140
THE CEREBRUM.
Laterally, it rests the superior lamina of the internal capsule, which separates against it from the lentiform nucleus. The thalamus is shaped like an
egg, with the small
It
Extremities.
in a large
The
thalamus
is lost
group of
posterior end (Fig. 44) a large pillow-like prominence, the pulvinar, and beneath presents it, are two smaller swellings; the outer one, which forms the lowest
The
point of the thalamus, is the lateral geniculate body; the medial The two geniculate bodies constitute geniculate body is the other.
the metathalamus (Fig. 43).
Surfaces.
immediate
down
as the
sulcus hypothalamicus (Fig. 28). It is joined to the internal surface of the opposite thalamus by the massa intermedia. It is
bounded above by the medullary stria. The superior surface is composed of a thin lamina of longitudinal white fibers, derived from the optic tract and radiation, called the stratum zonale.
The superior surface of the thalamus is divided by an oblique groove, the chorioidal groove, corresponding in position to the border of the fornix, into two areas a medial and lateral (Fig.
44).
The medial
area
is
ventricle and the fornix. Internally, it is bounded by the medullary stria of the thalamus. Posteriorly, next the stria, is a tribounded, behind, by a transverse groove in angular depression
front of the corpora quadrigemina, and by a slight groove, the sulcus habenulae, externally. That depressed surface is called Beneath the the triangle of the habenula (trigonum habenulae).
triangle is
The
lateral area of the superior surface is seen in the floor of the lateral
ventricle.
It
(tuberculum
nucleus
of
presents an anterior elevation, the anterior tubercle anterior thalami}, beneath which is the anterior
the
thalamus.
The
chorioid
sheet
of
epithelium,
extending from the fornix to the stria terminalis, covers this outer
141
Fig. 41.
143
area and separates it from the ventricular cavity. A lamina of fibers, the external medullary lamina, forms the lateral surface of the thalamus and rests upon the superior lamina of the internal
capsule.
Its fibers are
The
inferior surface blends with the superior surface of the tegmentum and substantia nigra, and forms the laminae and nuclei of the
tegmental hypothalamic region.
(See below.)
(Figs.
30 and 42).
The
(i)
of three layers:
and
(3)
inferior.
The nucleus
and
lies
hypothalamicus (Luysi)
ventro-lateral in
position
between the base of the internal capsule and the zona incerta. Like the substantia nigra just below it, it is composed of pigmented
gray matter. The reticular formation of the tegmentum, conThe tinuing beneath the thalamus, forms the zona incerta.
stratum dorsale
is
made up
as follows:
(a)
medial longitudinal bundle (Meynert); (b) the brachium conjunctivum (Forel), in which is the upper end of the red nucleus
of the
tegmentum; and
(c)
the medial
fillet,
which runs
lateral
and
The
It
lateral geniculate
body
43) forms a
of the lateral root of the optic tract the terminal nucleus of eighty per cent, of its fibers. It is joined to the superior quadrigeminal eminence by the brachium
and
superius.
In appearance
it
is
its
cell-bodies, is
divided into thick layers by thin laminae of fibers from the optic tract and radiation. The processes of the multipolar cell-bodies
It is geniculate body, placed at the end of the medial root, as the lateral geniculate is at the end of the outer root, of the optic tract. It rises up from
144
THE CEREBRUM.
is
and
brachium
front.
The medial
Its
laminated.
The brachium superius sweeps around it in geniculate body is gray in color and is not cell-bodies are small, and fusiform in shape.
They perhaps
give origin to the intercerebral fibers (Guddeni) and to a large part of the acustic radiation.
CEREBRUM.
SECTION
II.
THE MID-BRAIN.
cerebrum
the mid-brain
(MESENCEPHALON.)
The
44 and
third division of the
45).
It is situated
is
(Figs.
forms the connecting link between the fore-brain and the hindbrain. This has suggested the name "isthmus," sometimes
applied to
constriction
is
it:
though isthmus rhomb encephali refers only to the below the corpora quadrigemina. The mid-brain
vesicles, the
mesencepha-
lon (Figs. 16, 17 and 18). The cavity of the mesencephalon persists as the cerebral aqueduct, which is reduced to a slender canal by the thickening of its walls, roof and floor, due largely
to the
ingrowth of
fibers
of the brain.
The
cerebral hemispheres almost conceal the mid-brain from view; they overhang it dorsally, and the temporal lobes, inclosing it
its
anterior surface.
is
Only
the
median part
visible in the
complete brain (Fig. 21). The form of the mid-brain resembles a flattened cylinder. Its axis, a half inch long, is pointed upward and forward, and its long diameter, which varies from an inch
to
an inch and a
half in length,
is
directed transversely.
SURFACES.
The mid-brain has
which are
free,
four surfaces,
viz.,
the anterior
and
posterior,
and the superior and inferior, representing the ends of the cylinder, which are attached. The two latter are parallel with each other and are formed by section. The superior surface, sloping downward and forward, meets
SURFACES.
145
the anterior surface at an acute angle. Its inclination is that of the posterior end of the floor of the third ventricle. External to the floor of the ventricle, it is attached to the thalami and internal
capsules.
The
blending of
it
of the extreme lateral portions, the bases pedunculi, enter into In the median line the internal capsules of the hemispheres.
Fig. 42.
Transverse section of brain, cutting corpora mammillaria. (After Toldt, Morris's Anatomy.)
a. Lateral ventricle (central portion), b. Chorioid plexus of lateral ventricle, c. Caudate nucleus, d. Massa intermedia, e. Internal capsule. Lenticular nucleus: f, Putamen; gh. Zones, globus pallidus. i. External capsule, j. Claustrum. k. Ansa penduncularis. 1. Optic
tract,
m.
campus,
ventricle,
n. Inferior cornu of lateral ventricle, o. HippoOculomotor nerve, r. Corpus callosum. s. Fornix. t. Third Thalamo-mammillary fasciculus, w. Ansa lenticularis. x. Hy-
pothalamic nucleus (corpus Luysi). y. Substantia nigra. z. Basis of cerebral peduncle. bb. Interpeduncular fossa, cc. Pons (varolii.)
it is
The
little
superior surface
is
The
is
inferior surface joins the upper surface of the pons. It narrower than the superior surface. It is about one
inch and a quarter broad and measures an inch dorso-ventrally. The anterior surface (ventral) of the mid-brain looks forward
and downward
(Figs. 45,
47 and 48).
It is
146
udinally
slightly
THE CEREBRUM.
by a median sulcus, the fossa concave from above downward.
partially concealed
is
inter peduncularis,
It is
and
is
separated on either
side
lateralis mesencephali.
by the temporal lobes of the cerebrum, It is formed by a prominent basis pedunculi at either side; and by a median structure, band, the the posterior perforated substance which is inclosed between the
Though
two bases. The posterior perforated substance forms the floor of the median sulcus. The inner border of the basis pedunculi Thus is free and overhangs the perforated substance slightly.
is
formed the oculomotor groove (sulcus nervi oculomotorii) between the basis and perforated substance whence the third cerebral nerve takes its apparent origin. The fourth nerve courses forward
over the anterior surface, but
is
not attached to
it.
of the mid-brain
hemispheres.
cerebrum and
it
The
lateral sulcus
it
bounds
on each
side.
From
elevates
abruptly toward the median line, where it presents a longitudinal This produces two ridges which are subdivided by groove. a transverse groove into the four eminences, the colliculi of the
corpora
quadrigemina.
On
either
side,
is
anterior
and a
little
by an oblique
brachium
injerius.
The
ridges below the corpora quadrigemina are formed by the brachia conjunctiva of the cerebellum. The bottom of the
udinal
is formed by the superior medullary velum whence the trochlear nerve (fourth) is seen issuing.
I.
Mid-brain
<
TT
f
,
Tegmenta
,,
..
\
II.
Pedunculi
2.
Substantia nigra
3.
Bases pedunculi.
The four colliculi of the corpora quadrigemina and the four brachia connecting them with the geniculate bodies constitute
SURFACES.
147
the quadrigeminal lamina, which forms the greater part of the This lamina rests posterior surface of the mid-brain (Fig. 44). are
upon the dorsum of the pedunculi cerebri. The pedunculi cerebri made up of three great di visions, as shown above; named from
before backward, they are: (i) The bases pedunculi, comprising the [anterior part; (2) the substantia nigra, which is the middle
part;
and
(3)
The region of the mid-brain showing pulvinar of the thalamus, the genFig. 43. iculate bodies, the corpora quadrigemina and brachia, the pineal body, the
optic tract
a.
and
(Original.)
Chorioid groove, b. Medial geniculate body. c. Lateral geniculate body. d. Medial and lateral roots of optic tract, e. Optic tract, f Optic chiasma. g. Brachium inferius. h. Superior colliculus of corpora quad. i. Pineal body. j. Inferior colliculus of corpora quad. k. Brach. superius. 1. Thalamus. m. Fraenulumveli. n. Superior medullary velum. o. Fourth ventricle, p. Trochlear nerve.
.
lie in the posterior region next the quadrigeminal lamina. In the median plane between the quadrigeminal lamina and the 'tegmenta runs the cerebral aqueduct.
and
fibers, limited
and
half-inch
broad and
is
distinctly
longitudinally.
It
148
issues
THE CEREBRUM.
from the under surface of the fore-brain
at the junction of
the
median
hemisphere with the thalamus and, trending toward the At its superior end it is conline, descends to the pons.
tinuous with the motor tracts of the internal capsule. Four motor tracts make up the basis pedunculi, viz., the intermediate, the
the pyramidal, and the fronto-pontal. The deep portion of each basis pedunculi (Fig. 47) is occupied by the intermediate bundle, whose fibers rise in the corpus striatum
temporo-pontal,
in the nucleus pontis (Flechsig). The superficial should be studied in three parts. portion (1) The outer fifth of each basis contains the temporo-pontal tract (tractus cerebro-cortico-pontalis temporalis). It is composed
and terminate
which rise in the temporal cortex, in the superior, middle and inferior gyri (Dejerine); and, perhaps, in portions
of efferent fibers of the occipital lobe (Zacher)
and the
Pro-
ceeding through the inferior lamina and the occipital part of the
superior lamina of the internal capsule, and through the lateral part of the basis pedunculi, they terminate chiefly in the nucleus
of the pons; a
few end
in the
(Spitzka).
in the
The fibers are probably interrupted and relayed thalamus or lentiform nucleus. They form a segment of
the indirect motor path. These fibers are medullated later than the pyramidal tract (Flechsig).
(2)
The middle
is
and 48)
of the basis pedunculi (Figs. 47 occupied by the pyramidal tract (fasciculus longituthree- fifths
Its fibers rise in the anterior central
dinalis pyramidalis}.
gyrus
two-thirds of the occipital middle three-fifths of the basis, a part of the anterior longitudinal fibers of the pons, and the pyramid of the medulla. Below the
lateral
pyramidal
pyramidal tract which innervate the muscles of speech and of the face run through the genu of the internal capsule and constitute the medial portion
of the tract in the mid-brain
Those
fibers of the
(of
Bechterew). Immediately behind the face fibers, in the capsula interna, and external to them, in the basis pedunculi, are fibers
SURFACES.
149
Fig. 44.
The
dorsal or posterior aspect of the inter-brain, the mid-brain, the pons and the medulla. (Original.)
a. Anterior tubercle of thalamus. b. Pulyinar of thalamus. c. Brachium inferius. d. Inferior colliculus of corpora quad. e. Brachium conjunctivum. f. Corpus restiforme. g. i. Calamus scriptorius. h. Tuberculum acusticum. j. Area acustica. pontis. k. Posterior median fissure. m. Colliculus superior of corpora 1. Stalk of pineal body. quad. n. Medial geniculate body. o. Superior medullary velum, p. Median groove, q. Colliculus facialis. t. Trigonum n. hypoglossi. r. Fovea superior, s. Medullary striae, u. Fovea inferior, v. Ala cinerea. w. Taenia ventriculi quarti. x. Posterior lateral sulcus.
Brachium
SURFACES.
151
Still
of the
arm.
behind these, in
the internal capsule, and external to them, in the pyramidal tract of the basis pedunculi, are fibers for the innervation of the
trunk and leg muscles. (3) The inner fifth of the basis pedunculi is composed of the fronto-pontal tract (tractus cerebro-cortico-pontalis frontalis)
The origin of the latter is probably in the (Figs. 47 and 48). It is motor. This motor tract is contained prefrontal cortex. in the frontal part of the upper lamina of the internal capsule.
According to Hoche, it is interrupted and relayed either in the thalamus or striated body (Barker). Its termination is in the nucleus of the pons and in the motor nuclei of cerebral nerves
It constitutes a stage of an indirect motor path, like (Flechsig). the fibers of the outer fifth of the basis pedunculi, and the indirect
path
continued to the opposite half of the cerebellum by neurones whose cell-bodies are in the nucleus pontis (Flechsig).
is
The central part (Figs. 46, 47 and 48). a sheet of pigmented gray matter. The visible at the base of the brain between the
it
is
margin comes to
It
extends
to the corpora
Dorsal to
it
is slightly
concave longitudinally. The third nerve pieces it and comes out through the oculomotor groove. It contains small pigmented
multipolar cell-bodies, some of which constitute a relay for cerThere is a median tain fibers of the medial fillet (Barker).
cells located just in front of the pons, the ganglion (ganglion inter peduncular e). According to Forel, this ganglion is connected by a bundle of fibers, the fasciculus retroflexus, with the nucleus habenulae of the thalamus.
aggregation of these
inter peduncular
The
superior portion of the substantia nigra lies ventral to the nucleus hypothalamicus (Luysi) on either side. The nucleus hypothalamicus lies ventro-lateral to the red nucleus, and is
separated from
it
by the zona
incerta.
152
THE CEREBRUM.
(Figs. 46, 47
The Tegmenta
and
48).
The
posterior divisions
divisions, are in
a cover). They ventrally into the concavity bounded by the lateral sulcus
a triangle,
on the free side, where each tegmentum presents bounded by the sulcus lateralis, in front; by the braabove; and, inferiorly, by the lateral
fillet.
chium
inferius,
Dor-
sally, the tegmenta fuse with the quadrigeminal lamina except along the median line, where the cerebral aqueduct separates them.
Each tegmentum at the superior end blends with the thalamus, and helps to form the tegmental hypothalamic structures. Imbedded in that superior portion is the red nucleus (n. ruber) of
the tegmentum. (See tegmental hypothalamic region.) Inferiorly, the tegmenta are continued into the reticular formation of the pons.
The Cerebral Aqueduct (Aqueductus Cerebri, Sylvii, Figs. 17, 27 and 47). The aqueduct is a very slender canal connecting the third and fourth ventricles. So it is the "her a tertia ad quartum ventriculum." It is situated in the median line under the quadrigeminal lamina and produces a slight beveling of the dorsomedial borders of the tegmenta.
it is
It is
a half-inch long.
In shape
major and T-form, below, where it joins the fourth ventricle. Its height varies between a sixteenth and an eighth of an inch. Like other ventricles, it is lined with ependyma. A layer of gray
axis;
matter, thickest on the sides and floor, surrounds the aqueduct This is the stratum griseum centrale, which
some
The
stratum griseum centrale is continuous with the gray matter of In the ventral part of it are the nuclei of
the oculomotor (third), the trochlear (fourth) and the trigeminal (fifth) cerebral nerves.
Nuclei
48).
Oculomotor and Trochlear Nerves (Figs. 47 and Both nuclei together extend the entire length of the aqueof the
duct,
ventricle,
and the oculomotor is prolonged into the wall of the third where it receives a bundle of fibers from the opposite
SURFACES.
153
Fig. 45.
a.
(Original).
Interpeduncular fossa, b. Basis pedunculi. c. Pons. d. Trigeminal nerve, e. Abducent nerve, f. Acustic nerve, g. Facial nerve, h. intermediate nerve, i. Glossopharyngeal nerve, j. Vagus nerve, k. Accessory nerve. 1. Hyppglossal nerve, m. Anterior median fissure, n. Optic tract, o. Optic chiasma. p. Optic nerve, q. Stalk of infundibulum. r. Corpus mammillare. s. Oculomotor nerve, t. Medial and lateral roots of optic tract, u. Sulcus basilaris. v. Acustic nerve, w. Foramen caecum (Vicq d'Azyri). x. Pyramid, z. Anterior lateral sulcus. aa. Posterior lateral sulcus. bb. Funicy. Olive,
ulus lateralis.
SURFACES.
optic radiation
(n.
155
and optic
tract.
The nucleus
oculomotor ius), according to Perlia, is composed of seven distinct cell-groups, scattered from the posterior part of the lateral wall of the third ventricle down to the level of the transverse
groove between the superior and inferior quadrigeminal colliculi. From these seven cell-groups the axones proceed forward
into the nerve of the
same
median group
of cell-bodies
side; but there is, in addition, one which sends axones into both nerves.
The
by decussating
dendrites.
The
and substantia
The nucleus of the nigra and issue from the oculomotor groove. fourth nerve (n. trochlearis] is a single oval mass of cell-bodies
situated ventral to the inferior colliculus of the corpora quadrigemina. The root fibers of the fourth nerve, trochlear, proceed
dorsally and caudalward from the nucleus. They decussate with the fibers from the opposite nucleus in the superior medullary velum, from which they emerge on either side of the frenulum.
apparent origin.
The nucleus
is
composed
mid-brain
stratum griseum centrale, from the highest down to the pons. There is no break befifth
tween
this
nerve
formed by the substantia ferruginea under the locus caeruleus. The axones of these large cell-bodies run downward through
the nucleus, accumulating gradually until they form a distinct crescentic strand, which joins the chief motor root of the same
side.
The
homotemporo-pontal and
intermediate tracts) bring these nuclei into relation with the cerebral cortex; and the anterior and the medial longitudinal
Formatio Reticularis
(Fig. 47).
Through
156
THE CEREBRUM.
many oblique fibers interwoven with tracts of longitudinal fibers so as to produce a reticulum or net. Imbedded in the reticular formation ventral to each superior
quadrigeminal colliculus is a mass of large cell-bodies constituting the nucleus lateralis superior. This is the first of the reticular nuclei and the only one in the mid-brain. Many of the oblique
fibers of the formatio reticularis pass
tegmentum; they produce the tegmental decusat three successive levels, viz., the
rhomb
quadrigeminal colliculus (the fountain decussations) are: (i) The dorsal tegmental decussation (Meynerti) through which the anterior longitudinal bundle crosses. It is situated between the red
nuclei but dorsal to them.
(2)
The middle
(3)
the decussation
The ventral
teg-
mental decussation
47)-
(Foreli) in
which the
tract
At
is
the decussa-
tion of the
tivi).
It crosses
brachium conjunctivum (decussatio brachii conjuncwith its mate to reach the opposite red nucleus.
of
At
the
level
the isthmus
is
missure,
composed
of fibers
Intermingled with the above bundles of crossing fibers there are many other fibers of the formatio reticularis.
Tracts of Fibers in the Tegmentum (Figs. 46, 47 and 48). In the reticulum of the tegmentum there are eight distinct bundles of longitudinal fibers, viz., the anterior and the medial longitudinal bundles, the
fillet,
The medial
SURFACES.
157
running along the median raphe just ventral to the central gray In Weigert-Pal sections it shows clearly as a dark substance.
triangle nearly one-sixteenth of
will
an inch on a
side.
The
tract
be found in the same relative position in the pons and medulla oblongata. In addition to several very small strands 'of fibers
which
will
be explained
later,
is
Fig. 46.
Transverse section through the corpora mammillaria and the superior colliculi of the corpora (Original.) quadrigemina.
a. Lateral geniculate body. d. Brachium b. Thalamus. c. Medial geniculate body. superius. e. Pineal body. f. Stratum griseum centrale. g. Superior colliculus of corpora h. Formatio reticularis. i. Substantia nigra. k. Medial and Basis pedunculi. j. lateral nuclei of corpus mammillare. 1. Ventral tegmental decussation (Foreli). m. Dorsal tegmental decussation (Meynerti). n. Medial longitudinal bundle, o. Optic tract, p. Red nucleus, q. Medial fillet.
quad.
functionally
composed
of
two
tracts:
(i)
An
ascending bundle;
and
The ascending part of the medial longitudinal bundle is composed of fibers derived from the gray matter of the spinal
(i)
cord and from the terminal nuclei of sensory cerebral nerves. It is the continuation of the long ascending fibers of the anterior
fasciculus proprius.
Possibly a small portion of the tract runs through the posterior commissure to the thalamus and is common sensory in function but the major part of it decussates in several
;
158
THE CEREBRUM:
successive strands which end in the opposite motor nuclei of the The function of this latter part is reflex. cerebral nerves. (2) The descending part of the medial longitudinal bundle
is
composed
chiefly
of
cell-
Beginning
at the
nucleus lateralis superior in the mid-brain, it receives fibers from each reticular nucleus down to the nucleus lateralis inferior
of the medulla.
It receives
lateralis
the largest accession of fibers in the medius and the three nuclei
On
this
account James
tract.
S. Collier
suggests
that
it
It
through the anterior fasciculus proprius to the lower part of the Its size is gradually reduced by the ending of a few spinal cord.
fibers in the
cord.
fibers derived
In the mid-brain the medial longitudinal bundle also contains (i) from the oculomotor nucleus, which descend
to the pons,
and enter
the muscles of expression above the eye; and (2) from the nucleus of the abducent nerve. Running upward and decussating, the latter strand of fibers terminates in the opposite nucleus of the
oculomotor nerve, and thus innervates the internal rectus of that This strand accounts for the conjugate action of the two eye.
eyes in both health and disease.
distinct strand
and
It
48).
It
its
and
is
a descending
tract.
takes
whence
decussates at
once through the dorsal tegmental decussation (Meynerti) and descends through the reticular formation of the pons and medulla
and!, then,
through the fissural side of the anterior column of the Only by spinal cord until it fades away in the lumbar region. its degeneration has it been located. Its fibers end in the gray
matter of the cord on both sides and in the genetic nuclei of cerebral nerves; but, chiefly, in the nuclei of the oculomotor, trochlear and abducent nerves and in the cilio-spinal center of the spinal
SURFACES.
cord.
it is
159
connected with
retinae,
In function the anterior longitudinal bundle is reflex; all ocular reflexes which are excited by impulses such
as,
from the
accommodation
contraction and
dilatation.
The
laris,
Fillet or
Lemniscus
(Figs. 46, 47
and
48).
Near the
Fig. 47.
Section of the mid-brain through superior colliculi and the apparent origin of the oculomotor nerve. (Original.)
a.
Red
nucleus,
c.
g.
motor nucleus, e. Stratum griseum centrale. f. Colliculus superior of corpora quadrigemina. Formatio reticularis. h. Medial fillet, i. Medial geniculate body. j. Optic tract, k. Basis pedunculi. 1. Dorsal tegmental decussation (Meynerti). m. Ventral tegmental decussation
(Foreli).
n.
d.
Oculo-
r.
Pyramidal
Fossa interpeduncularis. o. Substantia nigra. p. Fronto-pontal tract, s. Intermediate tract, t. Temporo-pontal tract.
tract, q. ad.
N.
on
median raphe.
The
fillet is
into the ventral portion of the tegmentum, but immediately divides into two fasciculi, viz., the medial fillet, and the lateral fillet.
medial
Farther forward a small bundle leaves the lateral part of the fillet and runs up to the superior quadrigeminal colliculus.
That bundle
is
fillet.
l6o
Function.
tract.
THE CEREBRUM.
The
fillet
It carries spinal
muscular senses to
forms a segment in the direct sensory and cerebral impulses of the tactile and the corpora quadrigemina and thalamus,
to the inferior quadrigeminal colliculus.
f
L. Medialis
Lemniscus
-{
L. Superior.
<
{
L. Laterahs
,.
(Lemniscus medialis, Figs. 47 and 48). The fibers composing the medial fillet rise chiefly in the nucleus funiculi gracilis and nucleus funiculi cuneati of the opposite side of the
Medial
Fillet.
medulla oblongata.
lateral
They
cross
over in the
fillet
decussation
terminal nuclei of sensory cerebral nerves which cross the median plane and enter the opposite fillet. Thus connected with all common sensory nerves, and with the vestibular nerve, it enters
small bundle of the mid-brain and divides into two parts. fibers separating from the lateral part and running to the superior
quadrigeminal colliculus, forms the superior fillet. It associates movements with sensations from cerebral and spinal nerves. The medial fillet continues to the lateral nucleus of the thalamus,
ocular
tactile
sense.
fillet
From
to the
the thalamus the impulses are carried somaesthetic area of the cortex.
by the
cortical
lateral fillet (lemniscus lateralis) forms an oblique on the lateral border of the tegmentum (Fig. 44). It trends ridge upward and inward over the brachium conjunctivum to the inferior
The
few
quadrigeminal colliculus where some of its fibers terminate. fibers continue to the superior colliculus. Its function is
It rises chiefly from the ventral and lateral auditory conduction. of the cochlear nucleus (principally the opposite one) and parts ends in the inferior quadrigeminal colliculus. The greater
number
lary
its fibers cross through the trapezoid body and medulsome are uncrossed. It undergoes partial relay in the nucleus of the superior olive and nucleus of the trapezoid body on both sides and the nucleus of the lateral fillet on the same side.
of
striae,
The
lateral
fillet
is
SURFACES.
liculus
161
of the
into
the
brachium
proceeds from the inferior colliculus through the brachium inferius to the medial geniculate body and then through the acustic radiation to the temporal cortex.
Thus
the lateral
fillet
forms the
Fig. 48.
Section of the mid-brain cutting the inferior cofliculi of the corpora quadrigemina. (Original.)
a. Sulcus lateralis. b. Fonnatio reticularis. c. Medial longitudinal bundle, d. Nucleus of colliculus inferior, e. Aqueductus cerebri. f. Rubro-spinal tract, g. Lateral fillet, h. Medial filliet. i. Basis pedunculi. j. Location of anterior longitudinal bundle, k. Interpeduncular fossa. 1. Substantia nigra. m. Decussation of brachia conjunctiva.
conduction path. The acustic the lateral fillet the second stage, stage, the brachium inferius the third, and the acustic radiation the
second
stage
in
the
auditory
first
fourth stage. The last stage ends in the cortex of the superior and the transverse temporal gyri.
The spino-thalamic
of fibers not isolated
tract
(Figs.
47 and 48)
It is
is
located hi
a loose strand
has
many
relays in
course.
The
in the gray substance of the spinal cord and in the terminal nuclei of the common sensory cerebral nerves. Though it sends some
162
fibers to the
THE CEREBRUM.
quadrigeminal bodies, the substantia nigra and the lentiform nucleus, its chief termination is in the lateral nucleus
of the thalamus.
It
is
sensory.
It
conducts
tactile,
pain and
temperature impressions.
Brachium Conjunctivum. (Superior cerebellar The brachium conjunct! vum forms a ridge on the
quadrigeminal colliculus (Fig. 44). inward over its upper extremity. It
of the brachia conjunctiva
the median line of the isthmus, which ends above at the inferior
The
is
lateral
fillet
winds
by a
fibers
The
bend ventrally beneath the inferior colliculus of the corpora quadrigemina and, for the most part, decussate anterior to the cerebral aqueduct, through the median raphe (Fig. 48). These crossed fibers with the few uncrossed run forward toward the inferior surface of the thalamus, where
they inclose the red nucleus, and help to form the stratum dorsale of the hypothalamic region (Forel). Many of the fibers terminate
in the red nucleus
to the thalamus.
and from
it
Though most of the brachium conjunctivum rises in the cerebellum and forms a segment of an indirect sensory tract, it also contains efferent fibers which rise in the red nucleus.
Rubro-spinal tract (Crossed Descending Tract of the Red Formed by axones of the red nucleus, it immediately crosses through the ventral tegmental decussation (foreli) and proceeds lateral ward to the lateral fillet (Fig. 47). In the lower part of the mid-brain, it is imbedded in the medial part of that fillet. The rubro-spinal tract runs through the medulla and
Nucleus).
it
ends in
Olivary Fasciculus.
fibers
The
olivary bundle
is
a loose "strand of
to
traversing the
reticular formation
lateral
the medial
longitudinal bundle, in the upper part of the mid-brain; in the lower region of the mid-brain, it runs closer to the median line
and
is
fibers of the
brachium conjunctivum as
The
olivary bundle
probably rises in the lentiform nucleus and ends in the olivary nucleus of the medulla.
SURFACES.
163
of the
therefore,
It
occupies a thin crescentic area, just at the lateral border of the stratum griseum centrale, which thickens as
it
a motor root.
proceeds
downward toward
the pons.
it is
made up
few
fibers
in the
when
It is
upper part and only assumes a distinct shape and outline the level of the inferior quadrigeminal colliculus is reached.
continued to the middle of the pons in the same lateral relaand medial to the brachium conjunccerebelli
it
tivum
joins the
main
and bends
quadrigemina}.
The
groove
called
and
The
rior colliculus
and
is
circular in outline.
It
its
medial half the pineal body. It is joined to the lateral geniculate body by a band of fibers almost entirely concealed by the pulvinar
of the thalamus.
That band
is
is
the
superior colliculus
(Figs.
made up
It is
46 and 47).
composed
the stratum zonale, and a thick laminated gray layer, the stratum griseum. Within the stratum griseum many fibers end; a few from the lateral fillet, all of the superior fillet, and nearly all of the
brachium superius.
gives
origin
to
the
few
fibers that
It
an
optic-reflex center.
The
colliculus inferior of the corpora quadrigemina is elongIt is joined to the medial geniculate
ridge,
body by an oblique
called the
brachium
inferius,
it
and
it
from below,
164
THE CEREBRUM.
to the lateral
fillet,
due
cer-
ebellum.
Its surface is
made up
fillet
and brachium
which forms
the stratum zonale; gray substance, called the nucleus of the inferior colliculus, constitutes its deep portion. This nucleus receives
fillet
and gives
(Figs.
rise to
ium
inferius;
it
Brachium Superius
eral geniculate body with the superior quadrigeminal colliculus; but, excepting its extreme posterior end, it is buried in the substance of the thalamus. Its course is anterior and internal to
The brachium
relayed in the lateral geniculate body; it probably contains, also, a few optic fibers from the lateral root of the optic tract and a
few from the stratum griseum of the superior quadrigeminal colliculus. Optic reflex impulses are conducted by the brachium
superius.
Brachium
The
inferior arm,
sides.
Two
sets of fibers
the lateral
inferioris.
fillet
fibers
make up the brachium inferius, viz., and the axones from the' nucleus colliculi
in the medial geniculate body.
third
CEREBRUM.
SECTION
III.
ITS
We have noticed in the mid-brain that the basis pedunculi and much of the tegmentum are white matter, while the substantia nigra and quadrigeminal colliculi are composed chiefly of gray
substance.
all
of the inter-brain.
The
deep
165
Fig. 49.
human
nervous system.
(After Morris's Anatomy.) A. From spinal ganglion. B. From ventral horn of spinal cord. C. Pyramidal cell from cerebral cortex. D. Purkinje cell from cerebellar cortex. E. Golgi cell of type II from spinal cord. 'F. Fusiform cell from cerebral cortex. G. Sympathetic, a, axone; d, dendrites; c, collateral branches; ad, apical dendrites; bd, basal dendrites; cc, central process; p, perpheral process, the dendrite.
167
A thin envelope of gray the gray corpus striatum. substance, called the cortex, forms the surface of the hemisphere. The entire cerebral substance, both the white and the gray, is
which
is
composed
of neurones
and sustentacular
tissue,
made up of neuroglia, ependyma and mesoblastic connective tissue. Of necessity the cerebrum possesses a rich blood supply. The Neurone (Fig. 49). The essential element in the nervous system is the neurone. The neurone comprises the cell-body, its processes and end-organs. The cell-body, perikaryon, or
neurone
center, is
It
variable form.
a granular reticulated mass of protoplasm of ranges in man between four microns and one
hundred and
cell-body
Its shape,
is
thirty- five
microns in diameter.
The
size of the
which
onal,
dependent upon the number and mode of origin of its The cell-body has a large vesicular nucleus in which processes.
is
more
nucleoli.
It also possesses
a centrosome in
ganglia.
fibrilla
Its
reticulated
appearance
is
which form a
seen in
all
nucleus.
These
nbrillae
may be
from it.
Masses
produce the granulated aspect of the perikaryon. The tigroid bodies are found in the receptive side of the neurone, viz., in the
dendrites
cell-body,
except in the
axone-
in
axones.
position
protoplasmic in com-
and
where
they are smooth, like axones. In all cases they contain fibrillae prolonged from the cell-body. They branch richly and end in beaded points. The terminal branches are called telodendria.
may form synapses with axones of other neurones, terminate in special end-organs, or end free among the cells of any tissue. They are developed later than the axones and
Dendrites
may
An
1 68
THE CEREBRUM.
If
drites.
long
they
become
medullated.
Their conduction
is
cellulipetal, or afferent. Axones, neuraxones or axis-cylinders (Fig. 50) are smooth and fibrillar in character. One (occasionally two or more) is given off from the axone-
It
may
at once,
constituting a dendraxom; but usually it runs a considerable distance and sends out side branches, called collaterals. When
its function develops, and the medullary sheath presents segmental divisions separated by constrictions, called the nodes of Ranvier, at which points the colit
long, as a rule,
becomes medullated as
Both the parent axone and all its collaterals end by multiple division in the form of an end-brush, or tassel. This tassel lies in contact with the dendrites or body of another
neurone, with a secreting cell, or with a muscle fiber. In the last instance the fibers of the tassel spread out into disc-like platelets,
(Fig.
50).
Axone conduction
is
from the cell-body, cellulifugal, or efferent. The myelin sheath of axones and dendrites, which is developed as the neurone begins to functionate, is imbedded in neuroglia
cord
within the optic nerve and tracts, and in the brain and spinal but, elsewhere, is surrounded by the neurolemma (Schwann)
;
and the -fibrous sheath of Henle. The fibers of the olfactory nerve and most sympathetic fibers are not medullated, but they possess
fibrous sheath. Near the cell-body and near the end-tuft the processes are naked, having neither the fibrous nor the medullary sheath.
the
Types of Neurones.
preserves
its
i.
The
it
first
identity,
though
may
many
collaterals.
Found
2.
in brain
(Deiters).
type has a short axone, breaking at once into branches of apparently equal importance, the dendraxone. Found
in
The second
3.
cerebrum and cerebellum (Golgi). The third type has two or more axones
as in the
first
i.
diaxone, triaxone,
polyaxone
Orders of Neurones.
The
first
in relation with the periphery, as spinal-ganglion and anterior columna neurones, and conducts from the periphery or to it.
169
Fig. 50.
Motor neurone.
(After Barker.}
a.h. Axone-hillock devoid of Nissl bodies, and showing fibrillation, ax. Axis cylinder or axone; this process near the cell -body, becomes surrounded by myelin. m., and a cellular sheath, the neurilemma, the latter not being an integral part of the neurone, c. Cytoplasm showing Nissl bodies and lighter ground substance, d. Protoplasmic processes (dendrites) n. Nucleus, n.R. Node of Ranvier. s.f. Side n'. Nucleolus. containing Nissl bodies, n. of n. Nucleus of neurilemma sheath, tel. Motor end plate or telodenfibrils, collaterals, drion. m. Striped muscle fiber. s.L. Segmentation of Lantermann.
171
The second
with neurone- of
first
order has cell-body or distal process in relation It conducts to a neurone of the first order.
order or conducts centrally from it. In like are neurones of the third, fourth, fifth order, etc.
manner
there
Functions of Neurones.
Efferent.
i.
Afferent.
2.
Associative.
3.
neurone
is
afferent in function
if
when
is
its
its
axone
peripherally directed,
;Jeurilemma
Nerve-cell
Dendrite
Terminal branches
A
Fig. 51.
B
(After Brubaker.)
An
efferent
A. Efferent neurone.
function
is
An
is prevented from functionating, a slow atrophic change, called degeneration of Nissl. undergoes Rapid Wallerian degeneration occurs in an axone or dendrite w hich
r
completely cut off from all other parts of the neurone. In all probability neurones differ in chemical constitution and in electric
is
172
status,
cise
THE CEREBRUM.
hence chemicals, electricity and diseases appear to exera selective power and affect certain neurones without in(Figs. 52
cell,
fluencing others.
Development
rivative of
is
and
128).
Every neurone
is
the de-
an
epiblastic
which
This germinal cell, in the neural tube, sends out a process from what was originally its deep end, and becomes a neuroblast. This process forms the
axone and,
later,
During development
Fig. 52.
The
circles, indifferent cells ; circles with dots, neuroglia cells; shaded cells, germinal cells; circles with cross, germinal cells in mitosis; black cells, nerve-cells.
the cell-body wanders more or less from its original position, and thus reaches its adult location. The neurones are in this manner
aggregated into nuclei and cortex. The germinal cells of the neural crest form bipolar neuroblasts, which send out a process from each extremity. The axone appears first, the dendrite later,
as in
The
resulting
neurones
make up
the
cerebral, spinal
and sympathetic
and
in the
common
In the spinal ganglia ganglia. sensory cerebral ganglia, the neurones become
converted into the unipolar form by the lateral growth of the cellbody and the fusion, for a short distance, of the two processes
173
bipolar
cells
common stem
and
(Figs. 115
and
128).
They remain
in the olfactory
form multipolar
Sustentacular Tissue
(Fig. 53).
and
two forms
Fig- 53-
human
fetus, 23
cm. in length.
Showing the central canal with its substantia gelatinosa centralis, neuroglia Gordinier's Nervous System.) cells and ependyma cells. (After Lenhossek.
found supporting the neurones, viz.: (i) The comprising the neuroglia and the ependyma:
is
epiblastic tissue,
(A) Neuroglia most abundant in gray matter. It is made up of richly branched nucleated cells whose processes form a fine reticulation in the
174
larger
THE CEREBRUM.
meshes of the connective
tissue network.
and cerebrum, the processes of which are strong and richly branched; (b) the long rayed cells, located chiefly in the white substance, whose processes are long and fine and but little branched; and (c) the arborescent cells, which lie
in the cortex of cerebellum
Upon reaching the surface the branches form a limiting membrane of neuroglia. (B) The columnar ciliated ependymal cells which line the ventricles are also epiblastic in origin and should be classed with
the neuroglia cells. They form the only sustentacular tissue in the neural tube when it is first formed, at which time present
they extend from the ventricular to the exterior surface of the tube and, in the peripheral zone, form a rich supporting network. They appear to be of little importance in the adult condition.
(2)'
That
is
of mesoblastic origin
and
is
The
neurones constitute 53 per cent, of the brain and cord and the sUstentacular tissue 47 per cent.
(Donaldson). The white matter of the cerebro-spinal axis is made up chiefly of bundles of medullated axones imbedded in neuroglia and
supported
by connective
tissue.
The
fibers
possess
no
neuri-
lemma.
matter of the central nervous system is composed of cell-bodies and dendrites, chiefly, but also contains axones. These
The gray
nerve elements are supported by connective tissue and blood-vessels and are imbedded in a great abundance of neuroglia. The
nerve fibers in the gray matter are to a large extent non-medullated and naked.
is
con-
Ganglionar.
Central, or ventricular.
III.
175
P 11'
cgi
5 a
177
substantia corticalis consists of a thin envelope, the cortex (or bark), which forms the surface of the hemispheres and incloses
the white medulla, the centrum semiovale.
The
The
cortex varies in
thickness from a line to a quarter of an inch. Thickest on the surface of the gyrus, it grows thinner to the bottom of the sulci.
a reddish, or yellowish-gray color depending on the richness of the blood supply. By repeated observation of the symptoms
It is of
produced by
lower animal brains, both in the embryonic and adult condition, the cortex has been mapped out into certain definite functional areas (Figs. 54 and 55). Psychic
physiological study of
human and
function undoubtedly is dependent upon the associated activity of a number of cortical areas; but motor, common sensory and
special
sensory regions
exactness.
In
the following study of the cerebral cortex I shall use very exten-' sively the recent work of Dr. Alfred W. Campbell, entitled, " Histological Studies on the Localization of Cerebral Function," Cambridge, England. In this epoch-making work we are shown
that certain cortical areas have a characteristic histological struc-
ture that distinguishes them from all other areas. This will be referred to later under "cell and fiber lamination of the cortex,"
but to appreciate this histological evidence of localization one should thoroughly study the above work. Motor Area (Figs. 54, 55, 56 and 57). The emissive motor
area
is
and
in that part
it.
This
the center for ordinary voluntary motion on the opposite side of the body. Axones from this area descend to the nuclei of all
motor nerves.
area
In
lateral
sclerosis
there
is
degeneration and
motor
divided into four segments: the head (Campbell). and neck, the arm, the trunk, and the leg, named from below up-
ward.
The
first
178
THE CEREBRUM.
(Rolandi), though eye movements appear to be represented in the posterior end of the middle frontal gyrus; the arm area comprises the region between the genu inferius and the genu superius, the thumb, ringers, wrist, fore-arm, arm and shoulder move-
this ascending order; just above the shoulder area, at the genu superius, is the trunk area; and above that, in the anterior central gyrus and in the paracentral lobule,
The
the center for leg movements, representation in the leg center is inverted, the ascending
is
order being hip, thigh, leg, ankle, toes and great toe. The psychic motor areas, or areas for educated movements
are located just anterior to the above motor areas, in the anterior central gyrus and in the contiguous ends of the superior, middle
and
(Figs.
54 and 55).
psychic motor center for the lower extremity is located just in front of its emissive motor center in the probably anterior central and superior frontal gyri. In the posterior end
cortex.
is the psychic motor center for the the writing center of Gordinier; and in the inferior frontal arm, gyrus the center for the organs of voice and speech, hence the
The
cerebral hemisphere.
(Figs.
limited to the posterior wall of the sulcus centralis, including the anterior one-half of the posterior central gyrus and that part of the paracentral lobule which is continuous with it. This area
undergoes
(Campbell).
exclusive
It
is
Psychic Sensory Area (Figs. 54, 55, 56 and 57). large portion of the remainder of the parietal cortex probably constitutes a
number
of
centers
for
the interpretation of
common
sensory
impulses, hence the term, psychic sensor)' area. Impressions of the muscular sense are believed to be interpreted in the supra-
is
said to be located
179
lj
pilsi u o > u g o
E E
<u
-^^
Illpt
M "Q M "^3 g ~p
"*""*
l8l
and praecuneus. Perhaps other and temperature impulses. The parts interpret tactile, pain whole receptive and psychic area of common sensation has been
called the somasthetic area (Barker),
this
Acustic Center (Figs. 54 and 56). The receptive acustic center is located in the transverse temporal gyri and in that part of the superior temporal gyrus which is continuous with them. In the
left
adjacent part of the superior and middle temporal gyri, in the hemisphere, is the psychic acustic center.
Optic Center (Figs. 54 and 55). In the cuneus and lingual gyrus is located the receptive optic center for the temporal half
of the
same
retina
macula lutea
of both sides.
The remainder
of the
occipital lobe and, according to Mills and others, the angular This latter center is gyrus, also, form the psychic optic center.
left
hemisphere of
(Figs. 55
and
57).
The uncus
which
of smell, close to
in the fusiform gyrus is probably the gustatory center (Mills). Paul Flechsig in his recent studies of the human brain locates
taste
callosum
The
olfactory,
all
auditory,
visual,
common
distinguished by a definite characteristic histological structure, peculiar to each region (Campbell). Medullation of the fibers in these cortical areas occurs at different
times; and, according to Flechsig, in the following order: olfactory,
tactile
and muscular
and gustatory.
naming
center, the
center of equilibration,
and the
center of orientation.
nite
provided with projection fibers muscle groups and surface regions and with the organs of
and special sense areas are which connect them with defi-
182
sense.
THE CEREBRUM.
special
Large parts of the cerebral cortex possess no believed to be associative in function. projection fibers; they are
Association Centers of Flechsig.
Flechsig describes three asAnterior sociation centers, the anterior, middle, and posterior.
Association Center (Fig. 54).
of the frontal cortex
which
is
concentration, volition.
CONCRETE CONCEPT
Fig. 56.
Convex surface
of cerebral hemisphere.
(Brubaker's Physiology.)
(Fig. 56).
is
J. S. Bolton says of this association center that "it the last part of the cerebrum to be developed, and is the first
undergo dissolution; it is under-developed in amentia of all grades and atrophied in dementia, according to its degree." "It
to
The possesses the highest (mental) function" (Brain, Vol. 29). posterior association center, composed of those portions of cortex
situated
front,
between the sensory region of the equatorial zone, in and the visual cortex of the occipital lobe, behind, deter-
183
knowledge
cept"
association center.
(Fig.
To acquire thus the function of the posterior Mills calls it the center of "the concrete conis
It includes three psychic areas, the common 56). auditory and visual. Flechsig regards the island (of sensory, Lesions Reil) as the middle association center (Figs. 25 and 56). in it are associated with paraphasia.
Destructive lesions of parts of the motor or sensory cortex cause merely loss of certain motions and sensations represented
Fig- 57-
Medial and tentorial surface of Cortical areas after C. K. Mills. cerebral hemisphere. (Brubaker's Physiology.)
by those
parts,
the sensory, the psychic and the motor regions and causes aphasia, agraphia, change of temperament, impairment of the so-called
moral and
intellectual
faculties,
etc.
Ablation
of
the
visual
psychic center or auditory psychic center produces mind-blindness in the former and in the latter mind-deafness.
Cell
is
There
tions, prevails
a type of cerebral cortex which, with small but definite variathroughout the cerebrum, excepting in the visual
184
THE CEREBRUM.
(Fig. 58).
division
complicated,
It is to
it is
similar
I think
it
it
entirely
in this work.
and the cell layers have not been more satisfactorily this would assist in determining function. Dr.
gives the layers as follows:
:
Arthur
W. Campbell
2.
3.
4.
5.
6.
7.
cells.
2.
3.
4.
5.
6.
fiberless layer, or neuroglia zone. zonal layer, stratum zonale. supraradiary zone.
radiary zone.
felt-
work
of Kaes.
layer has next the surface a fiberless zone of
i.
The plexiform
neuroglia,
layer.
on account of which it is often called the neurogliaUnderneath the neuroglia is a more or less dense plexus
Scattered here and there in the zonal and supraradiary regions of the plexiform layer are small stellate cell-bodies, four or six
microns in diameter, belonging to the types of Golgi and Cajal, whose dendrites and whose double or triple axones ramify hi the
stratum zonale, some near the cell-body and others at a considerThe stratum zonale also contains dendritic able distance from it.
processes
Martinotti's
.*^
,
1 A
i -*
:
<
Cell and fiber lamination in the posterior half of the anterior central Fig. 58. The MOTOR AREA. (After A. W. Campbell's " Histological Studies on the gyrus. Localization of Cerebral Function." Published by the Syndics of the Cambridge
University Press.)
B. Stained to show only cell-bodies, z. Stratum zonale. A.. Stained to show only fibers. Supraradiary zone. B. Line of Baillarger. R. Radiary zone in the deep part of which the felt-work of Kaes. I. Plexiform layer. 2. Layer of small pyramids. 3. Layer of medium-sized pyramids. 4. External layer of large pyramids. 5. Stellate or polymorphous 6. Internal layer of large pyramids. 7. Layer of fusiform cells. cells.
S. is
187
from the commissural, the associative and the projection systems. It is very well marked in the motor area (Fig. 58), not so well in
the
sensory area (Fig. 59). In the uncus it is very distinct (Fig. 61) and is so thick and dense in the gyrus hippocampi The (the subiculum) as to be visible to the naked eye (Fig. 62).
zonal layer of
cortex.
fibers is faint in the visuo-sensory
common
and audito-
The stratum
zonalc
appears
to
grow
richer
The
iform layer
is
commonly thought
to
be association.
2. The layer of small pyramids (Figs. 58 and 59), as well as the third layer, is situated in the supraradiary zone. It is of small closely packed cell-bodies, pyramidal composed chiefly
in shape. They measure eight or ten microns in diameter. Their From the apices, surfaces and apices point toward the surface. lateral angles, dendrites are given off which ramify in the stratum zonale of the first layer. The axone issues from the base of the
pyramid and runs down through the subjacent layers. the small pyramids are a few ploymorphous cells.
3.
Among
a nearly
The
layer of medium-sized
pyramids
(Fig.
58)
is
uniform through-
is
In arrangement of cell-bodies and procThe pyramids get farther apart like the second layer.
size as the layer is descended. They measure ten to fifteen microns in their vertical diameter. In the anterior
central gyrus these cells are motor. Layers "two" and "three" be combined in one as wras formerly the custom. might
4.
layer
External Layer of Large Pyramids (Figs. 58 and 59). coincides in position with the line of Baillarger.
This
The
pyramids are larger and farther apart than in the above layer, and show a considerable accession of Nissl bodies as compared
with the smaller pyramids.
to 3o/z,
They measure
most important
i$fi to
20;*
by 25^
of the
criteria in dividing
the brain surface into different histological territories" (Campbell). As these cells, in the anterior central gyrus, degenerate in amyotrophic lateral sclerosis, they are considered motor in function. The apical processes appear to reach the first layer and ramify
in the stratum zonale; the lateral
1 88
THE CEREBRUM.
within the line of Baillarger; the axone runs inward to the white substance. The external large pyramids are found in nearly all parts of the cortex. They are replaced by a layer of large
stellate
fissure.
cells
The
in the receptive visual cortex along the calcarine line of Baillarger in the calcarine cortex is so thick
its
and dense that Gennari and, later,- Vicq d'Azyr described it from naked eye appearance (Fig. 60). In the motor and common sensor) area the line can scarcely be made out while in the supe7
;
there are two Baillargic lines, the deep line coinThere are ciding in position with the internal large pyramids. some peculiar Golgi cells among these pyramids, wiiose axones
rior parietal
gyms
run horizontally and give off collaterals that form pericellular baskets around the pyramids (Johnston). They are associative
in function.
5. The layer of stellate or polymorphous cells presents great variation in different regions and is a valuable guide for cortical
localization.
where
it
It is best marked in the calcarine area (Fig. 60) and the fourth layer are associated with the greatly thickit
is
situated
marked
in the auditory
it
is
common sensory region of the posterior The cell-bodies of the stellate layer 59).
They
are richly branched."
measure
five to eight
arborize at once near the cell-body. The axones of a considerable number of cells extend horizontally within the layer, helping
form the line of Baillarger; while others run toward the surface and end in the overlying laminae.
to
6. Internal Layer of Large Pyramids (Fig. 59). This is the most important layer of the cortex for localization. It lies in the radiary zone of fibers and is present in almost every part of the
by
60 and 61).
pyramids are
intermingled with irregular cell-bodies of the Golgi type and with Martinotti cells. The giant pyramids (Betz), or ganglionic
189
':
3
:
_.
'>?
--,
:
,-
: :
.:.-'
':
^U$>V
'
''^ "/
;':
';
';;..;.-
"
*^,-'.-
''.-
'
''\
"
**
-'
' '
\r
=''
.-'; "
:
-^
'"'''
fr
;}.''
"'
*^
^%'
ill
7
Fig. 59.
Cell
and
(After A. Studies on the Localization of Cerebral Function." the Cambridge University Press.)
gyms.
W. Campbell's
"
Histological
A. Stained to show fibers. B. Showing only cell-bodies, z. Stratum zonale. s. Supraradiary zone. B. Line of Baillarger. R. Radiary zone. I. Plexiform layer. 2. Layer of small pyramids. 3. Layer of medium-sized pyramids. 4. External layer of large pyramids. 5. Layer of stellate cells. 6. Internal layer of large pyramids. 7. Layer of fusiform cells.
IQI
(Bevin Lewis), are "pyraform" in shape. They are loaded with Nissl bodies and give off one axone and many dendrites. The
of the
lateral
and basal dendrites ramify in the radiary zone; the dendrite summit runs straight out toward the surface and like the
in the stratum zonale
The
the
and becomes a projection, association or commissural fiber. No where giant pyramids characterize the motor cortex.
"
Neither do they have in any other region definite nest-like grouping seen in
the anterior central gyrus. They measure twenty-five by sixty microns in the leg area of that gyrus twenty by forty- five microns
;
in the
arm
by
thirty-five
by Campbell, 87.5 per cent, of these cells in the affected area were entirely destroyed and those remaining showed signs of degeneration. Both the internal and external large pyramids
were thus affected in the anterior central gyrus, so they are motor in that gyrus; but they were not degenerated in any other part
of the cortex.
7.
The fusiform
variation.
layer
(Figs.
58,
59,
60 and 61)
very
is
little
found
topo-
cortex.
It presents
spindle-shaped cell-bodies lie in the deep part of the radiary zone and in the felt-work of Kaes. The long axes of the spindles are perpendicular to the surface in the crown of a gyrus but are parallel with it in the fissural walls and
floor.
From
off.
are given
ciation.
The
The
felt-work of
function of the spindle cell is probably assoKaes is a rich plexus of fibers in which
the white and gray substance meet. It is produced by the interof the association, commissural and projection fibers. mingling Scattered here and there through all the layers of the cortex are
two atypical neurones, viz., the dendraxones of Golgi, which arborize very richly and are associative in function; and, second, the
inverted
pyramids
of
Martinotti.
Of
terminate near the cell-body; but the axone runs out to the first layer and, branching T-like, ramifies in the stratum zonale.
192
Martinotti's
in function.
cells,
THE CEREBRUM.
like those of Golgi,
The Radiations
fiber zones
The six of Meynert (Figs. 58, 59, 60 and 61). above enumerated have been as fully indicated in
speaking of the cell lamination as the limits of this work will allow, but certain fibers, called the fibers of Meynert, need to be mentioned. The radiations of Meynert are strands of fibers,
clearly visible in the radiary
corticifugal
and
corticipetal
fibers.
The
corticifugal fibers
are
very largely axones from the pyramids; the incoming fibers of Meynert's radiations rise in other parts of the cortex or in gray
The radiating fibers belong to level. the following systems, the projection, the association and the commissural. The radiations are distinct as far out as the external
matter situated at a lower
layer of large pyramids and the line of Baillarger, hence the name radiary zone; but in certain regions they are much longer and in some parts of the temporal lobe they extend to the stratum
zonale of the
In such regions the line of Baillarger divides the radiary zone into two parts and there is no proper
first layer.
supraradiary zone. Association Fibers of Meynert. This name is applied to arcuate fibers of large diameter located in the radiary, the Baillargic
and the deep part of the supraradiary zone. Owing to condensation they form a distinct layer in the fissural walls and floor, but
crown of the gyrus (Campbell). arch over the medullary projection, crossing Meynert's They radiations at right angles. The deeper ones appear to be conare sparsely scattered in the
(Kaes and Campbell). Meynert's assoby Vulpius not to be developed before the
seventeenth year; and, according to Kaes, are most abundant in the highly developed parts of the brain.
Atypical Cortex.
The
and olfactory
areas.
The
and gyrus
cortex:
i.
marked variations from typical The greatly accentuated line of Baillarger, which
193
RECEPTIVE VISUAL Cell and fiber lamination in the calcarine region. Fig. 60. AREA. (After A. W. Campbell's " Histological Studies on the Localization of Cerebral Function." Published by the Syndics of the Cambridge University Press.)
A. Shows fibers of occipital cortex. B. Cells of same. z. Stratum zonale. s. Supra-radiary zone. G. Line of Baillarger or Genari. R. Radiary zone. I. Plexiform layer. 2. Layer of small pyramids. 3. Layer of medium-sized pyramids. 4. External layer of large stellate cells. 5. Small stellate cells. 6. Layer of giant pyramidal or stellate cells with some small
pyramids.
7.
Layer
of fusiform cells
IQ5
may be
was
first
seen dividing the cortex into two gray layers. This line seen and described in the visual area by Gennari and,
2.
The
is
replaced by a layer of
cells
give off three or four strong processes which appear to arborize in the line of Baillarger. They are found in the receptive and psychic visual areas but not elsewhere in the
cerebral cortex.
3. The internal layer of large pyramids is* rea conglomerate made up of three varieties of pyramids. placed by From without inward there are: First, small inverted pyramids,
cells
of Martinotti,
which extend
toward the
Second, the scattered giant pyramids, 25-30/1 in diameter, arranged in a single row and called the solitary cells of Meynert. The axones of the giant pyramids probably pass into
surface.
Olfactory Cortex.
are here mentioned, though only a part of them need be described the olfactory bulb, triangle, parolfactory area, anterior perforated
gyms
supra-
(longitudinal
striae),
fasciola
cinerea,
fascia
dentata,
hippocampus,
part of the
and uncus. The posterior inferior hippocampal gyrus and the gyrus cinguli are ordinarily
subiculum
included in the rhinencephalon; but, according to Elliot Smith, they belong to the neopallium (Figs. 26, 28 and 62).
The
is
layers as pictured
by Barker.
of the
These
named from
The
stratum
T-branched
fibers
These
fibers
with the surface for some distance, then bend centrally and break up into their end-tufts in the second layer. (2) The stratum
glomerulosum
are
is
made up
of
composed
brush-like dendrites from the spindle and mitral cells of the third and fourth layers. The glomeruli constitute the synapses between
the
first
ulare.
This
and second olfactory neurones. (3) The stratum reticis a network of mitral dendrites interwoven with
196
THE CEREBRUM.
fifth
layer
and the
cells.
brush
The
way
spindle cells likewise, both large and small, throw their dendritic processes down into the stratum glomeru-
second layer.
The
losum, where they end in rich tufts or brushes; and their axones penetrate the fourth and fifth layers, enter into the white sheath
of the bulb
(4)
stratum cellulare, or layer of mitral cell-bodies. The mitral cells have large pyramidal bodies with one axone and rich
dendritic processes. The latter arborize through the reticular to the glomeruli of the second layer, where they terminate layer
in the
The
form
of end-brushes.
The axones
run
colulti-
which they give off and then turn backward in the white sheath and
The
in the embyro.
derived from the ependymal lining of the ventricle (5) The stratum granulosum is composed of a
thick
layer of small cell-bodies, "granules," whose processes arborize richly in the granular, cellular and reticular layers.
in the granular layer are the medullated axones coursing
tract.
Imbedded
The
function
not understood.
cells of the olfactory bulb, it terminal nucleus of the olfactory nerves: the points of contact between them are established in the glomeruli; and the axones
The
and
its stria,
which
The lateral join the bulb to the cerebral hemisphere (Fig. 63). stria of the olfactory tract runs directly to the uncus, hence we shall study that region next.
The uncus and crown
of the
hippocampal gyrus
(Figs. 55, 6 1
and 62) probably represent the greater part of the lobus pyraformis of osmatic mammals. It constitutes the chief cortical
center of smell.
However
it
is
hippocampus,
fascia dentata,
and anterior end of the gyrus cinguli area of smell, as all showed arrested develop-
ment
in
197
Hill!
Fig. 61.
Cell
and
(After A.
W. Campbell's
Cerebral Function.
uncus hippocampi (lobus pyraformis). " Histological Studies on the LoPublished by the Syndics of the Cambridge
University Press.)
A. Showing fibers. z. Stratum zonale, external medullary B. Showing cell-bodies, lamina, s. B. Line of Baillarger. R. Supraradiary zone containing radiating filbers. Radiary zone. I. Plexiform layer. 2. Layer of stellate cells showing cell-nests. 3. Represents third and fourth layers of typical cortex, medium-sized pyramids obliquely placed and stellate cells; with Golgi's silver method shows tassel -cells. 4. Fusiform or triangular cells.
5.
Medium-pyramids.
6.
Fusiform
cells.
199
The
fascia dentata
is
of
first
importance accord-
Alexander
Hill.
He
Narwhal, which has no sense of smell, possesses every part of the hippocampal region excepting the dentate fascia (Campbell). The imcus comprises the whole anterior part of the gyrus hippocampi.
cells,
In structure the crown of the hippocampal gyrus and They have only five layers of
(i) As already pointed out the plexiform layer is thick and possesses a dense stratum zonale, only second to that of the subiculum. (2) The place of the small pyramids is usurped by the "olfactory islets" (Calleja) which are curious nests of large stellate cells (28^) interspersed with small nests of very minute
pyramidal
cells.
(3)
The
tassel-cells of Cajal.
Peculiar pyram-
such rich dendritic arborizations hanging from the bases as to resemble tassels, are seen in the place of the mediumidal cells, with
sized pyramids.
At the
typical
no
cells;
the
fourth
layer
of
cortex
is
entirely
wanting.
cells
(4)
The
r
stellate layer
of large
by a layer
of intermixed fusiform
and triangular
is
heavy w ith
Nissl bodies.
The
fusiform-cell layer
nearly typical.
Nucleus Amygdala (Fig. 32). In the anterior wall of the inferior horn of the lateral ventricle, near the temporal pole and
dorsal to the uncus hippocampi,
is
doubtful
classification.
The amygdala
in
part
continuous
with the corpus striatum and, according to Campbell, appears on the surface of the uncus as the gyrus semilunaris. Cortex of the Lower Wall and Lip of the Hippocampal Fissure (Fig. 62).
This
its
is
known
as the subiculum.
It is especially
distinguished for
remarkable stratum zonale, which is visible to the naked eye, and for its long radiations, which reach the
zonal layer and give the cortex a striated appearance, (i) The plexiform layer is almost wholly occupied by the stratum zonale, called here the external medullary lamina. (2) The layer of
olfactory islets.
The
islets
triangular cells, 5^ in diameter, resembling those in the uncus. (3) The stratum radiatum occupies about three-fourths of the
depth of
this
cortex.
In
its
200
THE CEREBRUM,
there are several layers of medium-sized pyramids, arranged in columns. The prominent apical processes of these pyramids
collect in
separating the columns of pyramids and producing the striations above mentioned. As the apical dendrites approach the olfactory
islets
The axones
of the
The pyramids
interruption through hippocampus into The alveus, which forms the the nucleus of the dentate fascia.
ventricular surface of the hippocampus, is made up largely of the axones of these pyramids; from the alveus they proceed into the crus of the fornix. (4) A few fusiform or stellate cells lie next
to .the type of Golgi, the axone being In function they are associative. It is wonderfully branched. in the region of these associative neurones that the axones of the
continue
without
the
the alveus.
They belong
pyramids bend and adjust themselves so as to enter the alveus nearly parallel with its surface, hence the name stratum oriens
applied to
it
by Edinger.
is
The
ture,
a free
lip of cortex
folded inward
It presents
a type of struc-
which
is
and the
marked
variation.
The stratum
zonale
not so prominent as
is
in the subiculum;
entirely replaced
by the nucleus
fasciae dentatae.
The nucleus
is
mids, of polymorphous and fusiform cells Their dendrites radiate toward the stratum zonale, their axones
The
dentate fascia
absent
Trigonum Olfactorium, Area Parolfactoria (Broca}, Gyrus Subcallosus and Septum Pellucidum, and Substantia Perjorata
Anterior (Figs. 26 and 28). These are the parts into which run most of the fibers of the medial and intermediate striae of the
olfactory tract.
They
are
more conspicuous
in the
embryo than
2OI
Fig. 62.
(After Edinger.)
203
is
human
brain.
The
The
plexiform layer
is
may
be
identified.
beneath that
occupied by separated by strands of fibers belonging to the olfactory tract and, perhaps, to the cingulum.
Central core of
scattered pyramids of
ependymal
cells
Fibers to C:
tract.
5.
S.
granulosum
4. S.
cellulare
3. S.
reticulare
2.
S.
glomerulosurn
i.
S.
nervosum
Nasal mucous
membrane
Olfactory cellbodies
Fig. 63.
(Gordinier after
Van
Gehuchten.}
The
an
is
characterized by
irregular direction of the pyramids and color affinity possessed by the deep cells.
cell layers,
and
and large cells, by an oblique by a most remarkable There are only four
(i)
The
plexiform presents a faint stratum zonale, (2) The layer of small pyramids
204
is ill
THE CEREBRUM.
defined.
(3)
layer of medium-sized pyramids placed at various angles occupies the place of the third, fourth, fifth and
sixth layers of typical cortex.
(4)
The
layer of spindle
cells.
In the spindle-cell layer are found the remarkable chromophilous cells. They are triangular or pyramidal in shape and have greater
affinity for stains
than the
cells of
cortex.
The claustrum (Figs. 31 and 42) is a sheet of peculiar gray substance which, according to Meynert, may be classed as cortical. In structure it resembles the seventh layer of typical cortex,
being
made up
of
fusiform cell-bodies.
The claustrum
is
The
external capsule is smooth, but the external surface is convoluted to coincide with the gyri insulae. At its lower border it joins the
lentiform nucleus.
II.
The substantia grisea ganglionaris is found in the great ganglia which, in their situation and relations, have already been considered.
They should be
They
are as follows:
1.
In the hemisphere:
The
2.
corpus
striatum,
composed
of the
le"ntiform nucleus.
In the inter-brain:
The
3.
thalamus, lateral and medial geniculate bodies, nucleus hypothalamicus (Luysi) and red nucleus.
superior and inferior quadrigeminal colliculi, the subnigra, and the nucleus lateralis superior in the
In the mid-brain:
The
stantia
tegmentum.
(Figs. 64, 31
and 66)
is
an ovoid mass
reddish-gray matter containing pigmented multipolar cellbodies of various sizes, those of large size being more numerous
in the nucleus lentiformis than in the nucleus caudatus.
The
205
Fig. 64.
Horizontal section of cerebrum through genu and below splenium of corpus callosum. Fornix and chorioid tela turned back, to show inter-brain
third ventricle.
and
a.
(Original.}
medullaris thalami (or pineal stria.) c. Chorioid Pineal body. f. Tail of caudate nucleus, g. Tapetum. Inferior longitudinal fasciculus, j. Anterior horn of of fornix. 1. Recessus triangularis. m. Anterior commissure.
b. Stria
e.
i.
Head
of caudate nucleus,
lateral ventricle, k. Columna n. Massa intermedia (or middle commissure), o. Posterior rigeminal colliculus. q. Posterior horn of lateral ventricle.
commissure,
p.
Superior[quad-
2O7
axones of those cell-bodies run both toward the pons and toward The corpus striatum, therefore, forms a
an
perhaps a
less
Of
the centrifugal fibers note the following three groups (i) The cortico-striate radiation made up of corticifugal and
:
Caudate nucleus
Stria terminalis
Pulvinar
Brachium superius
Inferior
quadrigeminal
colliculus
Medial geniculate
body body
Lateral geniculate
Corpus mammillare
Optic tract
Lateral stria of
oltactory tract
Olfactory bulb
Fig. 65.
Dissection of brain to
amygdalae,
etc.
corticipetal
fibers
the hemisphere. part crossed fibers, pass through the inferior lamina of the internal capsule and terminate in many nuclei of the inter-brain,
running from and to the equatorial zone of (2) The striato-thalamic fibers, which are in
mid-brain and pons; some of them pierce the superior lamina of the capsule. They run to the thalamus, hypothalamus and
208
THE CEREBRUM.
hypothalamic nucleus of both sides, the crossing fibers forming the superior commissure (Meynert's) in the tuber cinereum; to the
nucleus ruber and
quadrigeminal
colliculi;
and,
through the
deep part of the basis pedunculi, to the substantia nigra and nucleus pontis (Fig. 70). The last tract is the stratum intermedium,
or the intermediate bundle of the basis pedunculi (Fig. 68). (3) The olivary bundle (fasciculus olivaris) probably rises in the
t
and descends
is
probable
the corpus striatum or thalamus. The centripetal fibers connected with the corpus striatum are also numerous, (i) The thalamo-striate fibers, which belong
to the
sensory path, rise partly in the thalamus and They run through the frontal partly from nuclei lower down. stalk and the ventral stalk of the thalamus and, perhaps, through
stalk',
common
the parietal
fillet
fibers of the
medial
(2) Fibers from the (Fig. 70). to the posterior central cortex in the cortico-striate globus pallidus These radiation (the alpha and gamma bundles of Flechsig).
and spino-thalamic
up
in the superior
lamina of the
Lesions of the corpus striatum affect the internal capsule, which impales
it
;
and
may cause,
if
extensive, hemiplegia
and hemian-
66 and 31) is made up chiefly of gray matter containing multipolar and fusiform cell-bodies. The white matter consists of the stratum zonale on its free surface and
of the internal medullary lamina.
The thalamus
The latter divides the gray substance into nuclei, of which Nissl has described about twenty.
They may be grouped
as follows:
medial, lateral, anterior, nucleus of the pulvinar and nucleus habenulas. (i) The medial nucleus is joined to the opposite medial nucleus by the massa intermedia and is continuous with the hypothalamic
The
gray matter in the wall and floor of the third ventricle; but the
209
b.
hh
Fig. 66.'
Transverse section of the brain in the line of the pyramidal tracts, showing basal ganglia, internal capsules, corpus callosum, lateral and third ventricViewed from front. (Morris's Anatomy after Toldt.) les, etc.
c. Septum pellucidum. b. Radiation of corpus callosum. a. Longitudinal fissure, d. Chorioid plexus of lateral ventricle, e. Corona radiata. f. Column of fornix. g. Chorioid plexus of third ventricle, h. Internal capsule, i. Thalamus. j. Third ventricle, k. Interpeduncular fossa. 1. Inferior horn of lateral ventricle, m. Cerebral peduncle, n. Brachium o. Longitudinal pyramidal fasciculi of pons. p. Cerebellum, pontis. q. Deep fibers of t. Body of corpus callosum. r. Pyramid, s. Superior frontal gyrus. u. Anterior pons. horn of lateral ventricle, v. Head of caudate nucleus, w. Radiation of corpus striatum. aa. Claustrum. z. Insula. bb. Globus pallidus. cc. x. Putamen. y. External capsule, Optic tract, dd. Corpus mammillare. ee. Oculo-motor nerve, ff. Trigeminal nerve, gg. Facial and acoustic nerves, hh. Flocculus, ii. Glossopharyngeal nerve, jj. Vagus nerve. kk. Inferior olivary nucleus. 11. Decussation of pyramids.
211
the
of
same thalamus.
the
It gives origin to
thalamus.
rise to
some descending
of the cord.)
According to Bechterew it probably gives fibers that run down the cord. (Jas. S.
down
the lateral
column
(2)
The
It
which
fuse with
pulvinar.
it
posteriorly.
forms the terminal nucleus for the larger part of the tegmental fibers, especially of the medial fillet, the spino-thalamic tract, a part of the medial longitudinal bundle and the brachium conjunctivum of the cerebellum; and it constitutes the nucleus
It
of origin for
most of the
fillet.
Destruction
common
anesthesia and ataxia of the opposite side. (3) The nucleus of the anterior tubercle (Fig. 44) receives the fasciculus thalamo-mammillaris (Vicq d'Azyri) from the corpus
mammillare and
is
nix (Fig. 66). (4) The nucleus of the pulvinar (Fig. 44) is an important one. It receives about twenty per cent, of the optic fibers and gives
rise to
fibers in the
impairs vision.
(5)
It is
The
(Fig. 64).
fibers
nucleus of the habenula belongs to the epithalamus It receives It lies beneath the trigonum habenulae.
from the rhinencephalon through the medullary stria of the thalamus, and originates a bundle of fibers, the fasciculus retroflexus (Meynerti), which may be traced back through the
tegmentum to the interpeduncular ganglion in the substantia nigra. Beyond this, connections are probably established with the motor nuclei of cerebral nerves. The white matter of the thalamus includes, first, the stratum zonale of the superior surface, which is derived from the occipitothalamic radiation and the lateral root of the optic tract; and,
212
THE CEREBRUM.
second, the interior fibers, a part of which form the internal medulInto the thalamus enter the medial fillet, the spinolary lamina.
thalamic
tract,
a small part of the medial longitudinal bundle, cerebelli and perhaps some other
carrying
tegmental
fibers, all
common
whence the
proceeds
also
The thalamus
from the special sense paths, from the optic, auditory, olfactory, and probably the gustatory (?), and gives
receives
fibers
rise to fibers that
known
of
is
entered
by a considerable
number
through the occipitothalamic and temporo-thalamic radiations. The several bundles of thalamic fibers are as follows:
corticifugal
especially
(1) The columna of the fornix, having pierced the thalamus, descends to the corpus mammillare and terminates in its medial nucleus, whence the bundle of Vicq d'Azyr, the thalamo-mammil-
and ascends
to the thalamus.
It
ends in the
anterior nucleus.
(2) The stria medullaris thalami (Fig. 64) from the hippocampus and from the region of the olfactory triangle, terminates in the nucleus habenulse and from this nucleus the fasciculus retroflexus Both originates and descends to the interpeduncular ganglion.
to the olfactory paths. the lateral surface of the thalamus issue two groups (3) of fibers which rise chiefly in the lateral nucleus. They are
From
common
and
sensory in function:
(a)
The
inferior
rises
one
is
the ventral
which
lateral nuclei
it
and
is
passes toward its destination. It proceeds through the inferior lamina of the internal capsule and contributes fibers
to
under which
external capsule. Underneath the lentiform nucleus the ventral stalk is divided into two horizontal laminae by a thin sheet of gray
substance.
The upper
is
inter-
and enters
213
corpora quad.
Fig. 67.
The
optic path.
(Original.)
215
form nucleus, where it probably terminates; but the conduction path to which it belongs is continued to the upper third of the the paracentral gyri by the alpha and gamma posterior central and
bundles of Flechsig. The lower lamina of the ventral stalk, the inferior peduncle, runs principally into the external capsule medial
to the claustrum, but a part of
it
It
and the
The
superior
Fig. 68.
Section of the mid-brain through superior colliculi and the apparent origin of the oculomotor nerve. (Original.)
a.
Red
nucleus,
c.
d.
Oculo-
g.
motor nucleus, e. Stratum griseum centrale. f. Colliculus superior of corpora quadrigemina. Formatio reticularis. h. Medial fillet, i. Medial geniculate body. j. Optic tract, k. Basis pedunculi. 1. Dorsal tegmental decussation (Meynerti). m. Ventral tegmental decussation
(Foreli).
n.
r.
Pyramidal
Fossa interpeduncularis. o. Substantia nigra. p. Fronto-pontal tract, s. Intermediate tract, t. Temporo-pontal tract.
tract, q. ad.
N.
group of
from the
mus
is
It rises
from the
lateral nucleus
internal capsule.
To
is
gyms and
upper
in the paracentral
This parietal
stalk, in the
common
2l6
to the cortex.
THE CEREBRUM.
Flechsig has found that in development
it is
built
up
which receive
their
medullary
sheaths one after the other; and he names the bundles, according to the order of their medullation, after the first six letters of the
Greek alphabet
(4)
(see
page 232).
frontal stalk (Fig. 70) streams from the anterior end of the lateral nucleus via the frontal part of the internal capsule.
Its fibers
The
In
Fig. 69.
quadrigemina.
a.
(Original.)
Formatio reticularis. c. Medial longitudinal bundle, d. Nucleus of colliculus inferior, e. Aqueductus cerebri. f. Rubro-spinal tract, g. Lateral fillet, h. Medial filliet. i. Basis pedunculi. j. Location of anterior longitudinal bundle, k. InterpedunSulcus lateralis.
b.
cular fossa.
1.
Substantia nigra.
m. Decussation
of brachia conjunctiva.
The
all
ventral, parietal
and
the
common
fillet; the parietal stalk, high up, contains nearly sensory corticipetal fibers and, alone, is often
called cortical
fillet.
The
cortical
fillet
carries
common
sensations
from the medial fillet, the spino-thalamic tract, the medial longitudinal bundle and the brachium conjunctivum
received
21 7
up
Interruption of
the cortical
fillet
stops
all
common
Radiatio Occipito-thalamica (Gratioleti). A large pencil of fibers, the optic, or occipito-thaldmic radiation (Fig. 70) passes from the lateral geniculate body and the pulvinar through the internal capsule to the visual centers in the occipital lobe. A number of fibers in the occipito-thalamic radiation are corticifugal and
end in the superior quadrigeminal colliculus, though many are probably relayed in the thalamus and lateral geniculate body.
Destructive lesion of the occipito-thalamic radiation of either side produces hemianopia and atrophy of the same side of both retinae.
(6) Acustic
Radiation
or
Radiatio
radiates
Temporo-thalamica
(Fig.
70).
Another pencil of
fibers
thalamic radiation.
direction.
If the
area in the temporal lobe. It constitutes the temporoIts fibers are also in part corticifugal in
According to von Bechterew certain lesions in the thalamus have been accompanied by loss of facial movements expressing emotion. The red nucleus (nucleus ruber] of the tegmentum is situated
beneath the thalamus (Figs. 42 and 46).
in the indirect sensory tract,
It is
a relay-station
receiving the opposite brachium conjunct! vum cerebelli and, by its axones, continuing the tract to the thalamus and somaesthetic cortex. It also receives efferent
axones from the cerebral cortex (Beevor and Horsley) 'and gives origin to two bundles: (a) One centrifugal bundle of axones, (the rubro-spinal tract), after crossing over in the ventral decussation of the
tegmentum
(Forel's) descends,
;
first,
second, through the lateral area of the medulla, and, third, through the lateral part of the spinal cord. portion of the lateral
fillet
It
Gradually diminishing, it disappears at the first lumbar segment. ends in the lateral columna and center of the gray crescent of
(b)
The
axones through the opposite brachium conjunctivum of the cerebellum to the nucleus dentatus.
2l8
THE CEREBRUM.
(Luysi) (Figs. 30 and 42) is a bi-convex mass of gray matter placed ventro-lateral pigmented It to the red nucleus, and between it and the basis pedunculi.
is
It con-
an important relay for certain corticipetal fibers of the tegmentum and gives origin to others belonging to the cortical fillet. Certain descending fibers from the striate body terminate
stitutes
run through the tuber cinereum, just above the posterior border of the optic chiasma, and form the commissura superior (Meynerti), Gudden's commissure being
in this nucleus.
latter
The
called the
commissura
inferior.
is made up of two ganglia end of the thalamus, called the geniculate bodies at the posterior The lateral geniculate body (corpus geniculatum (Fig. 65). contains pigmented multipolar cells, which receive 80 laterale]
Metathalamus.
The metathalamus
per cent, of the optic fibers in the corresponding tract and give origin to a like per cent, of the corticipetal fibers in the occipito-
thalamic radiation; and, also, receive corticifugal fibers from the same radiation and originate a part of the brachium superius.
The
As the dark
cell-bodies are
appearance.
geniculatum mediate) is gray in color. Placed at the end of the medial root of the optic tract, it would at first appear to be concerned in vision; but it is not visual in function. Moreover, it is
The medial
optic
merely Gudden's commissure, the fibers of which are supposed to rise in the medial geniculate body. This body
contains chiefly fusiform cell-bodies which, in addition to their connection with the inferior commissure (of Gudden), receive
the end-tufts of fibers in the brachium inferius and send their
corpora quadrigemina. The latter are reflex in function. Ablation of the lateral geniculate body interrupts the visual
219
colliculi of the corpora quadrigemina (Figs. and 68) represent the optic lobes of birds, fishes and reptiles. 65
The superior
contain the center of optic reflexes. In being stratified, they bear some resemblance to the lateral geniculate bodies. They possess three white and two gray layers: (i) The stratum
They
zonale (stratum
album
superficiale) is
on the
This invests the laminated stratum griseum, which forms the deep part of the colliculus and comprises two
surface.
ficiale is
gray and two white layers: (2) The stratum griseum supercomposed of small multipolar cells. (3) The stratum album medium is a layer of fibers separating the small from the
cells.
large multipolar
(4)
The
large cells
make up
the stratum
griseum profundum, underneath which is another layer of fibers. The fibers of the superficial, (5) The stratum album profundum. middle and deep strata comprise, first, those that enter the colliculus through the optic tract
and
lateral
fillet,
spino-thalamic tract; and, second, those that take origin in the colliculus and leave it through the brachium superius or the
anterior longitudinal bundle.
Of
superior colliculus
it is
supposed that
and running through the brachium superius some go as far as the retina; probably others
fillet.
The
is
made up
of efferent axones
crosses at once
,It
through the dorsal tegmental decussation (Fig. 68) and descends ventro-lateral to the opposite medial longitudinal bundle, to the anterior columna of gray matter in the spinal cord. Its fibers
third, fourth
and
sixth cerebral
nerves and in the cervical enlargement of the spinal cord; but perhaps others enter the remaining nuclei of motor cerebral
nerves,
and a few fibers of the tract have been traced as low as lumbar region. This bundle is the great optic reflex tract. The fibers to the nuclei of the third, fourth and sixth cerebral nerves bring about the reflex movements of the eyeball, contraction
the
220
of the pupil
THE CEREBRUM.
and accommodation
to distance; while those fibers
which end in the gray substance of the lower part of the cervical
enlargement of the spinal cord, called the cilio-spinal center, through the white rami communicantes and cervical sympathetic,
The
Destructive lesions affecting the superior quadrigeminal colliculi produce loss of reflex movement of the eyeballs, loss of pupillary
reflex
and
loss of
accommodation.
The
quadrigemina form
a relay in the auditory path (Figs. 65 and 69). They are made up of a white stratum zonale, whose fibers are continuous chiefly
with the lateral
fillet
and brachium
inferius,
and
of a
deep gray
of small
colliculi iuferioris,
which
is
composed
multipolar cell-bodies in a
network of
fibers.
The
nuclei of the
considerable number of fibers belonging to both lateral fillets, but most of them belong to that of the same side; and from them proceed axones of the auditory paths through the brachia inferiora A few fibers of the spino-thalato the medial geniculate bodies.
mic
tract also
colliculus.
Again
this colliculus
Though
the
greater part of
the lateral
fillet
passes by the
apt to involve the entire bundle and cause almost complete deafness in the opposite ear.
body
is
Nucleus Lateralis Superior (Fig. 68). In the reticular formation of the tegmentum at the level of the superior quadrigeminal
colliculus
is
the
multipolar
cell-bodies.
It
contains
large
and descending fibers of the Tschermak, a small fasciculus runs from this nucleus According into the medial longitudinal bundle where it divides T-like; and its descending fibers run down through the anterior fasciculus
ascending
to
proprius of the cord (Barker). The spino-thalamic tract probably undergoes a partial relay in the superior lateral nucleus.
221
Substantia Nigra
68 and 69).
The
merited multipolar cell-bodies which make up nigra form, -first, a terminal nucleus for certain fibers of the medial fillet and a nucleus of origin for other fibers which continue in
that tract (Barker); and, second, a terminal station for the fasciculus retroflexus (Meynerti)
tract
Beyond
probably continued, but with the exception of the intermediate tract they have not been traced. The interefferent tracts are
mediate tract
III.
is
It is
the hypothalamus;
tricle,
the
(3)
The lamina
cinerea
and 27) form a sheet of gray substance that connects the inferior and medial surfaces of the cerebral hemispheres and may be called their inferior
terminalis
gray commissure. The optic chiasma is white matter, and the hypophysis is not composed of nerve tissue at all and, therefore, neither one need be described in this place. From the floor of
the third ventricle the gray matter extends laterally beneath the thalamus, and is continuous with the anterior perforated subof the floor also extends up to the sulcus on the medial surface of the thalamus. The hypothalamicus inferior gray commissure receives efferent fibers from the corpus striatum of both sides. Some of these fibers form a commissure
stance.
just
it
is
called the
commissura
inferior
The
Meynert's com-
(Figs. 26
and
46).
The
corpora mammillaria (albicantia), though composed of fornix fibers on the surface, contain in the interior two nuclei, the medial
and
lateral.
The medial
nucleus
is
It
222
receives the
THE CEREBRUM.
the fibers in the columna of the fornix end-tu^ts of to the fasciculus mammillaris princeps. The gives origin
and
sending one branch, the thalamo-mammillary bundle (of Vicq d'Azyr), up to the anterior nucleus of the thalamus and the other branch, the fasciculus pedunculo-mammillatter bifurcates,
laris,
backward into the tegmentum. The fasciculus thalamomammillaris (Vicq d'Azyri) connects the fornix with the thalamus. The tegmental part of the pedunculo-mammillary bundle has
been traced, in the mouse by Cajal, downward to the lower part of the pons, running anterior to the medial longitudinal bundle
(Barker).
The
bundle, which according to Flechsig ends in the substantia grisea centrale of the mid-brain and is thence connected with the motor
nerve-nuclei and the automatic centers of the medulla (Barker). Through the fornix, the stria medullaris thalami and the fascic-
ulus
retroflexus,
and through the fornix and the pedunculoreflex connections of the olfactory
nerve
(the
and 64)
when
It is formed, joins the medial nuclei of the thalami. present, by the approximation and fusion of the thalami
in the second
month of embryonic life. It is occasionally absent. In the massa intermedia are cell-bodies and transverse fibers.
The
It is
latter
line; at least
many
of the fibers
do not cross
(3)
not a commissure in the ordinary sense of that term. The stratum griseum centrale of the mid-brain (Figs.
and 69) surrounds the cerebral aqueduct (Sylvii). This matter begins in the lateral wall of the third ventricle. It gray extends through the mid-brain and is continuous with the gray
46, 68
and a part
con-
According
to
223
The
nucleus of the
is
an elongated
mass
centrale,
down
to the level of the transverse groove between the quadrigeminal colliculi. The nuclei are placed somewhat obliquely; at the lower end they fuse in the median plane. According to seven distinct cell-nests are found in each nucleus; and Perlia,
nest fifteen occupies the area of fusion and is common to both nerves. The greater number of axones of this nucleus run for-
ward
same side; but those from the median and a small bundle from each nucleus
descends with the medial longitudinal bundle to the colliculus facialis, where it joins the facial nerve and through that nerve supplies the muscles of facial expression above the orbit.
Trochlear Nucleus (Fig. 69).
is
The
corpora quadrigemina. It is in the ventral part of the stratum griseum centrale like the oculomotor nucleus. Unlike the third, the axones from the nucleus of the fourth cerebral
colliculus
the
nerve run backward and issue from the posterior surface of the mid-brain at the isthmus they are peculiar also, in that the axones
;
decussate before their emergence (Fig. 65). The nuclei of the oculomotor and trochlear nerves receive
fibers
other motor tracts of the internal capsule and thus obtain their
voluntary motor and inhibitory impulses. It is probable also that the third nucleus receives fibers, through the medial longitudinal bundle, from the opposite abducent nucleus, and that the part of the nucleus which receives these fibers supplies the For the purpose of reflex both internal rectus muscle of the eye. the oculomotor and trochlear nuclei receive fibers from the ante-
and medial longitudinal bundles, from the pedunculo-mammillary bundles (?) and, perhaps, from the cerebellum through the brachia conjunctiva.
rior
224
THE CEREBRUM.
is
fifth,
and
is
motor nucleus of the trigeminal. ing root of the fifth nerve, which descends
joins the
merely the superior end of the It gives origin to the descendto the
main motor
root.
In
its
course
downward
chium conjunctivum
cerebelli.
tegmentum. If so, the result is paralysis of the nerves on the same side and hemianaesthesia, hemiataxia, loss of taste (?) and
deafness on the opposite side. The white matter of the
cerebrum
is
composed, in the
adult condition, of medullated fibers; the medullation begins in is continued for a considerable
time after birth (Flechsig). Within the cortical substance the myelin sheaths continue to be laid down until late in life (Kaes, Mc-
Murrich).
1.
The
cerebral fibers
form three
definite systems:
2.
3.
Association fibers.
I.
PROJECTION FIBERS.
and and
projection fibers are connected only with the motor sensory areas of the cerebral cortex and are, therefore, motor
The
sensory in function (Figs. 54 and 55). Where they are present they are continuous with Meynert's radiations. They are composed, first, of the medullated axones of the pyramids and the
polymorphous neurones; these descend from the cerebral cortex, are motor in function, or corticifugal, and constitute the upper
motor segment and, second, they comprise the medullated axones whose cell-bodies are situated in gray matter below the cerebral cortex; these axones ascend to the cortex and are
;
of neurones
The
PROJECTION FIBERS.
22$
capsule and the mid-brain, and vice versa (Figs. 30 and 42). They connect the cortex, directly or indirectly, with all parts of the body, throwing or projecting a picture of every part and
Many
Within the hemisphere all projection fibers run through one great sheet, the internal capsule, with the exception of the olfactory; but in the mid-brain, they are separated into two great groups
the basis pedunculi and the tegmentum, the substantia nigra
intervening.
CORTICIFUGAL, OR
The most important
fibers are the following,
MOTOR PROJECTION
FIBERS.
motor projection the intermediate tract, the frontonamely, pontal tract, the pyramidal tract and the temporo-pontal tract.
tracts of corticifugal or
The intermediate
70,
69 and 87) extends from the corpus striatum through the inferior lamina of the capsule and the deep part of the basis pedunculi to the motor cerebral nuclei and to the nucleus pontis, though
it is
probably relayed in the substantia nigra. From the nucleus pontis axones run by way of the brachium pontis to the cortex
The
intermediate
efferent, or
motor path.
tract (tractus cerebro-cortico-pontalis jrontalis. Figs, 70, 71, 69 and 87) rises from the cortex of the frontal lobe anterior to the precentral sulci. It tranverses the centrum
The fronto-pontal
semiovale, corona radiata, frontal part of the internal capsule one-fifth of the basis pedunculi to the ventral area
of the pons,
where it terminates in the nucleus pontis (chiefly) the nuclei of motor cerebral nerves (Flechsig). It is probably relayed in the thalamus (Beevor and Horsley).
According to Dejerine, the temporo-pontal tract
bro-cortico-pontalis temporalis, Figs.
70, 71,
(tractus cere-
lamina (and posterior part of the superior lamina) of the internal capsule and lateral one-fifth of the basis pedunculi to the substantia nigra and the
inferior
15
226
THE CEREBRUM.
nucleus pontis; but according to Spitzka some of its fibers end in the nuclei of motor cerebral nerves. Thus it should be noted
with the exception of those fibers to motor nuclei of the cerebral nerves, each of the three tracts above mentioned, viz.,
that,
the intermediate; fronto-pontal and temporo-pontal, constitutes a segment of an indirect efferent path which is interrupted in the
nucleus pontis and then continued by the axones of that nucleus through the brachium pontis of the cerebellum. It is probable, though not surely established, that the fronto-pontal and temporo-
Fig. 70.
much
enlarged.
BLUE, Common sensory tracts: Fr. St., Frontal stalk; Par. Stalk, Parietal stalk; Ventral stalk. RED, Motor tracts: Fr.-P. Tract, Fronto-pontal tract; Pyr. Tract, Pyramidal tract; Temp. -P. Tr., Temporo-pontal tract Interm. Tract, Intermediate tract. PURPLE, Special sense tracts: Opt. R., Occipito-thalamic radiation; Acust. R., Temporo-thalamic
;
radiation.
pontal tracts are relayed in the corpus striatum or thalamus, as they have been found undegenerated in the base of the peduncle
when their cortical origins were destroyed by extensive lesions. The pyramidal tract (tractus cerebro-spinalis pyramidalis)
(Figs. 70
and 71)
from the giant pyramids and large polymorphous cells of that Descending through the corona radiata, genu and region.
anterior two-thirds of the occipital part of the internal capsule, the pyramidal tract comprises the middle three-fifths of the basis
PROJECTION FIBERS.
forms the pyramid of the medulla and the anterior and
pyramidal
tracts of the spinal cord (Figs. 71, 69, 87, 93
227
lateral
and
102).
with a few exceptions, cross over to the opposite side; they end in connection with the motor nuclei of cerebral and spinal nerves. Fibers enter the nucleus of
fibers of the
The
pyramidal
tract,
the trochlear (or fourth) nerve chiefly on the same side, and a few descend to the motor nuclei of other cerebral nerves and to the gray matter in the spinal cord without decussation; all other pyramidal fibers terminate on the side opposite to their origin. The fibers from the lower one-fourth of the anterior central gyrus,
which go
to the
motor nuclei
Fig. 71. Diagram of internal capsule in colors. (Original.) RED, motor; BLUE, common sensory; PURPLE, special sensory.
leave the pyramidal tract high up hi the peduncle and run for some distance through the medial portion of the fillet; they constitute Bechterew's accessory lemniscus.
This accessory
fillet
has
70, 105
and
106).
Those
fibers
the genu inferius of the central sulcus. They run through the genu of the internal capsule to the peduncle and, then, both through
228
THE CEREBRUM.
the accessory fillet and the inner portion of the middle threefifths of the basis pedunculi. Upper Extremity Fibers (Figs. 70
and
105).
The
fibers of the
and through
it
innervate the
muscles of the upper extremity, take their origin from that part of the anterior central gyrus adjacent to the foot of the middle
frontal gyrus: their origin lies
intersect
superius, re-
run through the pars occipitalis of the spectively. internal capsule just behind the genu, and through the basis pedunculi immediately lateral to the head and neck fibers. Those
fibers
fibers
rise
These
lowest
which innervate the muscles of the thumb, fingers and hand, down in the arm area of the cortex and occupy the
arm bundle
and the
The fibers which control the in the peduncle. shoulder muscles rise in the upper part of the cortical area and form the anterior and medial part of the arm bundle in the capand basis pedunculi, respectively; while the wrist, forearm, elbow and arm are innervated by means of fibers which Trunk Fibers. are intermediate in both origin and course.
sula interna
The trunk
fibers of the pyramidal tract rise in that projection of the anterior central gyrus which is situated just above the genu superius of the central sulcus. In the internal capsule, the trunk fibers
A large number of
and carry impulses to the nerves of the lower extremity. They originate in the upper fourth of the anterior central gyrus and in the paracentral lobule. The hip fibers rise farthest downward and the toe fibers farthest upward,
enlargement of the spinal cord
centralis.
The
fibers
have
same
base of
b, b. End-tufts of sensory fibers in cortex, c. Nucleus of funiculus cuneatus, showing end-tufts of fibers from the cord. d. Nucleus of funiculus gracilis, containing end-tufts of fibers from cord. e. Section of medulla at fillet decussation. f. Section of medulla at pyramidal decussation. g, g. Motorial end-plates. b. Section of cervical cord, showing terminations of fibers of anterior and lateral pyramidal tract, i, i. Spinal ganglia, j, k. Short sensory fibers. 1. Long sensory fibers, m, m, m. Sensory end-organs, n. Section of lumbar cord.
a, a.
Motor
PROJECTION FIBERS.
229
Fig. 72.
PROJECTION FIBERS.
231
the peduncle the hip fibers are medial and the toe fibers lateral. Fibers which innervate the muscles of the thigh, leg and small
toes
have
and great
this same relative position and order between the hip toe fibers both in their cortical origin and in their course
There are other corticifugal fibers in the internal capsule, viz., some within the occipito-thalamic and temporo-thalamic radiations (Figs. 70 and 71) and others running from the special and
common
sensory areas of the cortex; but these fibers are probably reflex- in function and do not properly belong to the projection group.
Several bundles of descending fibers are found in the tegmentum, namely, the anterior longitudinal bundle, which is reflex in func-
from the red nucleus, a small part of the brachium conjunctivum of the cerebellum, the descending root of the trigeminal nerve, the olivary bundle, and certain other
tion, the rubro-spinal tract
fibers
in
the formatio
is
reticularis.
With
these
exceptions the
in function.
tegmentum
ascending in direction
and sensory
Destruction by clot or tumor, or otherwise, of any of the above divisions of the pyramidal tract causes upper segment paralysis
of the particular muscles innervated through that tract, the muscles
The
mic
tract; the
of the formatio
the paths of conduction are continued through the internal capsule. The medial fillet carries impressions of the tactile and the
tract
conducts
tactile,
pain
and temperature impulses; while all varieties of common sensory impulses are carried by the brachium conjunctivum cerebelli.
Chiefly through these three tracts,
common
sensory impressions
232
THE CEREBRUM.
arrive in the lateral nucleus of the thalamus. In the capsula interna the corticipetal projection fibers constitute the cortical The fillet and the optic and acustic (and gustatory?} radiations. former end in the somaesthetic area of the cerebral cortex, the
latter in the visual, auditory
and gustatory
are
cortex.
The
olfactory
projection fibers
tegmentum nor
the
directly
The
not been entirely determined, but it is in the lateral nucleus of the thalamus.
thalamus (Fig. 70) runs through the internal capsule, in the inferior
lamina. It is relayed largely hi the globus pallidus. Its fibers enter the medullary laminae of the nucleus lentiformis and the external capsule; ultimately they terminate in the upper onethird of the posterior central gyrus and in the temporal and insular
lobes (see p. 212).
From
a great pencil of fibers, called the frontal stalk (Fig. 70). It mingles to a small extent with the fibers of the pyramidal tract, but runs
chiefly
Its
termination
some
of
its fibers
The parietal stalk issues from the lateral surface of the thalamus higher up than the ventral stalk and mingles with the pyramidal fibers in the superior
Its
location
is
It terminates in the para(Figs. 70 and 71). and the posterior central gyrus. According to Flechsig some fibers also end in the upper extremity of the anterior central, the superior frontal gyri and in the underlying part of
of the capsule
central lobule
the gyrus cinguli. The parietal stalk, if we may apply this name to all the common sensory fibers in the occipital part of the capsule, ultimately comprises all the corticipetal fibers to the somaesthetic
area.
The alpha
PROJECTION FIBERS.
lidus to the
central.
/3.
233
Origin
in doubt.
large bundle
nucleus to the
central gyrus
7.
upper
It runs from the globus large. same area as beta bundle. Origin is in doubt. 8. Runs from the lateral nucleus of thalamus (from the central of Luys and the arcuate nuclei) to the middle third of the central
pallidus to
lateral nucleus
to the
f.
lower third of the posterior central gyrus. The zeta bundle is last to be medullated.
It rises in
the
superior part of the lateral nucleus and terminates in the foot that (posterior end) of the superior frontal gyrus and in
part of the gyrus cinguli which underlies the foot of the
frontal.
first
The cortical fillet conveys common sensory impulses to the If the cortical fillet be somaesthetic area of the cerebral cortex.
severed,
all
common
interrupted; and complete loss of common sensation on the opposite side of the body and hemiataxia result. Just where the fibers conducting taste impulses are located in the tegmentum is unknown Auditory impulses run through
fillet and the brachium inferius to the medial geniculate while optic impulses run directly to the lateral geniculate body; body and the pulvinar without passing through the mid-brain
the lateral
at
all.
at present
Within the internal capsule the gustatory tract cannot be located; but the acustic and visual paths are well
or temporo-thalamic radiation (Figs. 70
known.
The
acustic,
and 54)
and the
third
and fourth
is
fifths of
the
superior temporal gyrus (Barker). produces deafness in the opposite ear, which
not complete
is
234
corticifugal
fibers
THE CEREBRUM.
in the temporo-thalamic
probably
reflex in function.
70, 38, 45
The
and
body and
thalamus.
It
retrolentiform region of the internal capsule to the cortex of the Half -blindness in the same side of lingual and cuneate gyri.
both
retinae
results
from section
of the
optic
radiation.
The
by Campbell
to
be axones of the
to
corpora quadrigemina, where they end in contact with the neurones of the anterior longitudinal bundle. Their function is
reflex.
II.
COMMISSURAL FIBERS.
opposite sides of the cerebrum and, like the projection fibers, are continuous with the radiations of Meynert. They are contained chiefly in the corpus callosum, the
They connect
and the commissura hippocampi; but are found in the posterior commissure, commissura hebenularum, inferior (Gudden's) and superior (Meynert's) commissures.
anterior commissure,
also
as already described,
is
between the cerebral hemispheres (Figs. 34, 27 and 73). Its fibers connect both similar and dissimilar parts of the cortices; within the hemisphere, they form a prominent radiation, called
The corpus callosum is made up almost wholly of cortical axones, a few of them being projection fibers; of a small number of afferent projection fibers, and of
the radiatio corporis callosi.
collaterals
fibers,
from the association and projection fibers. All callosal except the few efferent projection fibers, end on the opposite
It is the
corpus callosum,
makes
it
cerebrum
to act together as
of recent development,
since
COMMISSURAL FIBERS.
235
opposite temporal and occipital lobes together (pars occipitotemporalis], the limbic lobes with the contra-lateral olfactory
tracts,
tracts with
oljactoria).
It is
not joined by the great commissure, especially the cortex of the tentorial areas of the cerebral hemispheres. In size it varies
Fig-
73-
Transverse
section
of
Viewed from
front.
Commissural
fibers.
Anatomy
after Toldt.)
a. Caudate nucleus (head), b. Internal capsule (frontal portion). Lentiform nucleus: c. Putamen, d Glpbus pallidus. e. Medullary lamina, f. External capsule, g. Claustrum. h. Vena terminalis. i. Interventricular foramen (Monroi). j. Anterior perforated substance, k. Uncus. 1. Anterior commissure, m. Longitudinal fissure, n. Corpus callosum. o. Anterior horn of lateral ventricle, q. Septum p. Chorioid plexus of lateral ventricle, pellucidum. r. Columns of fornix. s. Lateral fissure (Sylvii). t. Gyri of insula. u. Optic recess, v. Optic tract, w. Optic chiasma. x. Inferior commissure (Guddeni)
.
inversely as the corpus callosum. Its importance diminishes with the appearance of the corpus callosum in the lower mammalia
and
it
Below mammals it is said to be the most important connecting link between the hemispheres and is philogenetically very old.
236
THE CEREBRUM.
the lyre (Fig. 36), unites the
hippocampal gyrus, dentate fascia, and the hippocampus with This is the commissure of their fellows of the opposite side.
the pyraform lobes, the cortical areas of smell.
III.
ASSOCIATION FIBERS.
These fibers remain on the same side and connect parts of the same hemisphere. They are situated within or beneath the
cortex, the various parts of
fibers
effort
is
to unite.
Association
become medullated and actively functional only as mental and education gradually develop them. So far as the brain
first,
They unite
more numerous and are contiguous parts of the same gyrus and
In direction
they comprise arcuate and tangential fibers and they are intracortical and subcortical, in position. Every zone of the cerebral cortex
contains association fibers, from the felt-work of
Kaes
to the
stratum zonale.
zone and adjacent part of the supraradiary zone, along the line
of Baillarger (Fig. 60);
and (2) in the zonal layer (Figs. 58 and Those sparsely scattered large fibers whose location is 61). (i) indicated by the line of Baillarger are called Meynerfs association
fibers.
The deeper
if
of these fibers are continuous with the radiato the short asso-
tions of
ciation fibers,
the
more
and
are truly associative in function. The associative fibers of Meynert are compacted together by pressure in the walls and floor of the sulci. In the crown of a gyrus they are scattered. Their
exact origins are not yet worked out; but they are probably the horizontal processes of cells in the second to sixth layers. (2) The association fibers of the plexiform layer of the cortex, which
constitute
(Fig.
61),
when
ASSOCIATION FIBERS.
237
contiguous parts within circumscribed areas. The richness of the zonal layer of fibers, as already pointed out in describing
the plexiform layer of the cortex, varies greatly in different regions, being best developed in the subiculum. The fibers comprising
the zonal layer
(a)
dendrites of the cells of Cajal in the plexiform layer, (b) The (c) The T-branched apical dendrites of the subjacent pyramids,
Fig. 74.
Diagram
of association fibers in the cerebral hemisphere. (Gordinier and Quain after Meynert.)
Short association fibers, connecting adjacent gyri. f.l.s. Fasciculus longitudinalis suc.i. Cingulum. f.p. Fasciculus perpendicularis. f.l.i. Fasciculus longitudinalis inv.d'A. Thalamo-mammilf.u. Fasciculus uncinatus. fo. Fornix. fi. Crus fornicis. lary bundle of Vicq d'Azyr.
s.
perior,
ferior,
axones of Martinotti's
cells,
(d)
The
corticipetal axones
which
terminate in the superficial layer of the cortex. The short association fibers are almost infinite in their connections.
They connect
and
the
their interruption
on the
the receptive and psychic sensory areas, left side causes inability to interpret
sterecalled mind-blindness, mind-deafness, Again, those short fibers also associate the psychic with the psychic-motor, and the psychic-motor with the emissive-
sensations,
agnosis, etc.
238
TRE CEREBRUM.
centers.
motor
In
this
manner
is
connected
with the motor center for the upper extremity, and the speech center with the motor centers for the lips, tongue, etc.: breaking of the former connection on the left side destroys ability to write,
agraphia; and aphasia results, if the latter connection is broken. Besides these and many other connections of associated centers,
the short fibers join together the various parts of each cortical
area.
into bundles.
The long association fibers (Figs. 74 and 75) are collected They rise from the pyramidal, the polymorphous
layers of the cerebral cortex (Cajal),
and are
Proceeding out of the lobe in which they rise, being interlobar, they dip down into the centrum semiovale and arborize
known
The
less distant parts of the cortex. Among are the following bundles: cingulum of the gyrus fornicatus (Fig. 74) is a bundle
which almost
perforated substance the gyrus cinguli and hippocampal gyrus, to the uncus through and temporal pole. The fibers, which form several systems,
callosum.
It
anterior
have been divided into three groups by Beevor, namely: (a) The anterior, which joins the region of the anterior perforated substance to the fore part of the frontal lobe, (b) The horizontal, which unites the frontal lobe and the gyrus fornicatus. (c) The
which associates the lingual and fusiform gyri with the hippocampal gyrus and the pole of the temporal lobe. Like the two following bundles it establishes associations
posterior fasciculus,
The Fornix
(Fig. 74).
In each
a bundle of association fibers as well as of projection and commissural fibers. The projection fibers rise in the olfactory bulb
and in the region of the olfactory triangle and, running up through septum pellucidum to the fornix, continue through it to their destination in the hippocampus and uncus. The commissural fibers of the fornix rise in the uncus, the fascia dentata and gyrus
hippocampi and run through the alveus of the hippocampus into the crus fornicis, whence they cross through the commissura hip-
ASSOCIATION FIBERS.
2 39
pocampi to the opposite side (Fig. 36). They terminate in the dentate fascia and hippocampal gyrus, including the uncus. The associative group of fibers in the jornix has the same origin as the
commissural group.
the
It enters
cms
fornicis
body
NCfT)
pCR.
NG
Fig. 75.
Stria terminalis and fasciculus uncinatus. Fasciculus occipito-frontalis. (Gordinier after Dejerine.)
Cge. Lat. geniculate body. Cgi. Med. geniculate body. coa. Anterior commissure. Fu. Fasciculus uncinatus. Gh. Ganglion of the habenula. NA. Amygdaloid nucleus. Na. Anterior nucleus of thalamus. NC. Head of caudate nucleus. NC'' Tail of caudate nucleus. NC(T). Body of caudate nucleus. OF. Fasciculus occipito-frontalis. OF (Tap). Part of the fasciculus occipito-frontalis forming the tapetum. pCR. Foot of corona radiata. Pul. Pulvinar. sch. Choriodal fissure. Tga. Columna of fornix. Th and Th(Va). Thalamus. tsc(lc). Stria terminalis. tth. Stria medullaris thalami. n. Optic tract.
it
The
smaller
is
the stria
medullaris thalami (pineal stria) (Fig. 40), which bends backward and runs along the supero-medial border of the thalamus to the
it
larum.
By the fasciculus retroflexus of Meynert, it with the inter-peduncular ganglion of the mid-brain.
continues
as
connected
The
larger
bundle
columna
fornicis
(Fig.
74).
As already
240
indicated,
it
THE CEREBRUM.
terminates in the medial nucleus of the corpus mamon the opposite side. Its termination is associated
millare, partly
first,
with the anterior nucleus of the thalamus by the thalamomamrnillary bundle (of Vicq d'Azyr) and, second, with the midbrain and pons by the pedunculo-mammillary bundles. (3) The uncinate fasciculus (fasciculus uncinatus, Fig. 74) is a bundle, with some sharply curved fibers, which arches over
the
main stem of the fissura cerebri lateralis, and connects the uncus and the anterior temporal region with the orbital gyri and
the pole of the frontal lobe. It is situated near the basal surface. Its fibers spread out at both ends in the cortex, and they especially join the
Like the cingulum and forgyri with the limbic lobe (Barker). Lesion in any one nix, it is connected with the rhinencephalon. of these three bundles causes disturbance of smell.
(4) The superior longitudinal fasciculus (fasciculus longitudinalis superior, Fig. 74) is a sagittal bundle located beneath
the convex surface of the hemisphere, just above the posterior ramus of the lateral fissure of the cerebrum. According to Cun-
ningham, it runs just above and behind the putamen of the lentiform nucleus, external to the base of the corona radiata. Its
fibers diverge at the posterior
end of the lateral fissure and radiate and temporal cortex: some of them,
The
run as far forward as the temporal superior longitudinal bundle joins the frontal cortex
fissure,
with the parietal, occipital and the external temporal. It thus associates the psychic auditory and visual centers with the motor
is
the result of
its
interrup-
(5) The inferior longitudinal fasciculus (fasciculus longitudinalis inferior, Fig. 74) is about on a level with the lateral ventricle. It passes near the outer wall of the inferior and posterior
horns of that ventricle, being separated from them by the occipitothalamic radiation and the tapetum. It connects the temporal lobe to the occipital. In the temporal lobe its fibers cross at
lamina of the internal capsule. This fasciculus unites the auditory and visual psychic centers, and
right angles those of the inferior
ASSOCIATION FIBERS.
241
thus associates the concepts of things seen with those of things heard. Breaking of this connection produces intercortical sensory aphasia.
gest
its
an object seen is recognized, but fails to sugname; and the spoken word is heard and understood, but
So,
it
repre-
sents.
to Ferrier
inferior longitudinal
is
present in the
The
Fasciculus
Occipito-frontalis
(Foreli).
This
is
fibers formerly
75).
It
is
between the callosum and the internal capsule, just external to the lateral ventricle. It extends from the cortex of every part of
the frontal lobe to the cortex of the convex surface and lateral
border of the occipital lobe. Posteriorly, the fibers diverge to form a fan-like sheet in which there is an intermingling of fibers from the corpus callosum (Cunningham); and that sheet enters
into the external
ventricle
boundary
of the inferior
and
horn, hence the synonym, tapetum. The tapetum is lined by the ventricular ependyma and is separated from the inferior longit-
Its particular
Perpendicularis,
a very broad vertical bundle located just in It extends from the inferior parietal front of the occipital pole.
Fig. 74).
and superior
down
to the
temporal, the inferior occipital and the fusiform gyri. Its function classed with the short association fibers.
doubtful.
CHAPTER
IV.
THE RHOMBENCEPHALON.
SECTION
I.
THE CEREBELLUM.
composed
is
is
It is the lozenge27). the dorsal and largest portion of it, and is developed in the roof-plate and dorsal zones of the metencephalon. Together with the pons it forms the hind-brain.
(Figs. 20, 21
and
The
cerebellum
weight is about five ounces, slightly more than one-tenth of the whole brain. It is situated in the posterior fossa of the skull,
Its
and dorsal to the pons and medulla oblongata. Between it and the last two structures is enclosed the fourth ventricle. The cerebellum is distinguished from the
under the tentorium
cerebelli
cerebrum by
its stratification. Its surface is composed of gray the cortex (substantia corticalis); its interior is white substance, and is called the medullary body (corpus medullare, Figs. 79 and 81). Function. The cerebellum is an important relay in the in-
direct
motor and indirect sensory paths. In response to impulses received from skin, muscles, tendons, joints and viscera, it is also
and
maintain
equilibrium.
through the brachia conjunctiva, upon the opposite side of the cerebrum.
Divisions.
The
cerebellum
is
made up
of
two
lateral parts,
the hemispheres, and a central part, uniting the hemispheres together, called the vermis cerebelli, or worm (Figs. 76, 77, and 80).
In the early embryo the cerebellum is a transverse ridge in the roof of the fourth ventricle, partially divided for a time by a median
groove on its ventricular surface; and it remains undifferentiated into medial and lateral parts in many lower animals (Edinger).
(hemispheria
cerebelli)
measure
THE CEREBELLUM.
243
two inches from before backward and about the same in thickness, antero-medially but they taper rapidly toward the lateral
;
borders (Figs. 76 and 77). They-are joined together by the worm, or vermis, which forms the central and most elevated part of .the
cerebellum.
The vermis
cerebelli, or
worm,
is
Fig. 76.
Superior surface
(Original.)
a. Pineal body. b. Colliculus superior of corp. quad. c. Lateral sulcus. d. Colliculus inferior of corp. quad. e. Culmen monticuli. f. Pars posterior of quadrangular lobule, g. Superior semilunar lobule, h. Anterior tubercle of thalamus. i. Stria medullaris thalami. k. Mid-brain. 1. Inferior horn of lateral ventricle, m. Pars anj. TrigonunvhabenulEe. terior of quadrangular lobule, n. Predeclivil sulcus. O. Postdeclivil sulcus. p. Declive monticuli. q. Folium vermis. r. Posterior cerebellar notch, s. Horizontal sulcus.
,
and much thinner than the hemisphere (Figs. 76 and 80). In animals lower than mammals, it is not differentiated from the hemispheres and appears to be the only part of the cerebellum
shorter
present, being very large in birds
Its transverse ridges give
it
and swimming
reptiles (Edinger).
It unites
a worm-like appearance.
the upper half of the medial aspect of the two hemispheres, their
244
THE RHOMBENCEPHALON.
separated
by an
antero-posterior
The upper
inferior.
vermis
is
worm, or vermis
superior;
and the
superior
The
and
terior
riorly,
end of the
worm by
at. the
the hemispheres, the anterior and posterior cerebellar notches. The posterior cerebellar notch, incisura cerebelli posterior
(Fig.
76),
bounded by the
worm and
the
falx
postero-medial border of the hemispheres, is occupied by the A prolongation of the medullary body of the cerebelli.
cerebellum
fills
up the incisura
cerebelli
anterior,
or anterior
cerebellar notch, which is situated between the antero-medial borders of the hemispheres in front of the vermis cerebelli.
is
the white
splits, in its
median
bands (peduncles), and an inferior, which is the inferior medullary velum (Figs. 79 and 78). Separating at an acute angle, the two laminae form the tent of the fourth ventricle.
pairs of connecting
The
is
inferior medullary
is
inferius, Figs.
79 and 91)
the inferior lamina of the medullary body. It a short plate of white matter, not more than a quarter of an
is
separated from the superior lamina by the angle, It ends in a concave border from which a
down
and together they form the inferior half of the roof of that cavity. Laterally, the inferior velum extends to the flocculus of the hemisphere.
It
Of
the
worm
it
BNA
"
v.
m. posterius;" but
I have used
is
velum medullare anterius" and no more reason for those embryological terms
"
inferius
"
in this place
than there
and
"superius" and
anatomy.
their
positions in adult
THE CEREBELLUM.
245
joins the cereis
of the
medullary body
The
It
superior lamina
(cerebellar
made
connecting bands
peduncles)
longations of the corpus medullare of the cerebellum, the inferior velum (Fig. 91).
except
Fig- 77a.
(Original.)
Tonsil, d. Superior medullary velum, e. Lobulus centralis. f. Culmen monticuli. g. Inferior medullary velum, h. Brachium conjunctivum. i. Restiform body. j. Brachium pontis. k. Peduncle of flocculus. 1. Division in biventral lobule, m. Lobulus gracilis. n. Lobulus biventer. o. Prepyramidal sulcus. p. Nodule, q. Uvula, r. Depression in tonsil, s. Postnodular sulcus.
Horizontal sulcus.
quadrigeminal colliculi, where they disappear. They are joined to one another by a thin plate of white matter, the superior medulWith the velum, they lary velum (velum medullare superius}.
lateral
They
boundaries of the superior half of the gradually bury themselves in the pons
upward toward the corpora quadrigemina. Beneath the corpora quadrigemina and the cerebral aqueduct, the brachia conjunctiva cerebelli decussate, and pass into the
246
THE RHOMBENCEPHALON.
hypothalamic region of the opposite side. They end chiefly which their fibers surround. Near the corpora quadrigemina each brachium conjunctivum is obliquely crossed
lateral fillet in its course to the inferior
by the
quadrigeminal
colliculus.
and
wider
it fuses with the corpus medullare of the cerebellum than mesencephalic end. It forms the floor of the groove between the brachia conjunctiva cerebelli and the superior half of the
where
at the
Fig. 78.
Dissection of rhombencephalon to
pontis
and corpus
restiforme. (Gordinier,
On left side the cerebellar brachia and restiform body have been cut short; the right is cut obliquely to show connection with brachium conjunctivum and corpus restiforme. i. Median groove of fourth ventricle. 2. Medullary striae. 3. Restiform body. 4. Clava in fuhiculus gracilis. 5,5. Brachium conjunctivum. 6. Lateral fillet. 7,7. Lateral sulcus of mid-brain. 8. Corpora quadrigemina.
hemisphere
Its lateral
unite the brachia conjunctiva, hence their name. In the median line its posterior surface presents a slight ridge, the fr&nulum
veil,
from
issue
either side of
from the cerebellum betwe.en the brachium conjunclum) tivum and the brachium pontis (Figs. 78 and 86). They first run
forward to the posterior surface of the pons, near the inferior
border; and then, bending
(a flexion
THE CEREBELLUM.
of
247
more than 90
floor
of the
form the
ventricle.
and
to
bound
pons (Figs. 78, 85 and 91). They are continuous with the transverse fibers in the ventral area of
to the lateral borders of the
Fig. 79.
a.
Median
and medulla.
(Original.)
Declive monticuli. d. Postdeclivil sulcus. e. h. Postpyramidal sulcus. Folium vermis. f. Horizontal sulcus. g. Tuber vermis. i. Pyramid, j. Prepyramidal sulcus. k. Uvula. 1. Culmen monticuli. m. Postcentral sulcus. n. Central lobule, o. Inferior colliculus of corp. quad. p. Cerebral aqueduct, q. Precentral s. Lingula. t. Medial longitudinal bundle, r. Superior medullary velum, sulcus. u. Fastigium. v. Inferior medullary velum, w. Nodule, x. Postnodular sulcus.
Predeclivil sulcus.
Arbor
vitae.
c.
the pons. The brachia pontis in the anterior cerebellar notch are placed external to the brachia conjunctiva and the restiform bodies, and are opposite the widest part of the fourth ventricle.
The
Horizontal Sulcus of Cerebellum (Figs. 76, 79 and 80). cerebellum has one great sulcus which divides it into upper
surface.
and lower
The
is
irregularly
body
248
THE RHOMBENCEPHALON.
dividing the border of each hemisphere and the posterior end of the worm. Rarely the two halves are not continuous through the
posterior extremity of the
worm.
remaining important
sulci
They
are nearly parallel with one another; hence, the cerebellum is laminated, not convoluted like the cerebrum. Though the
is an important landmark in the adult cerebellum, does not form a primary embryonic division of the cerebellum but appears late in foetal life (Cunningham).
horizontal sulcus
it
bounded by the horizontal sulcus and the superior lamina of the medullary body (Figs. 76 and 79). The posterior and larger part of this surface is covered by the tentorium cerebelli, the tentorial area; the small anterior part of it bounds the anterior
The superior surface is divided into rive continuous lobes by four crescentic sulci, called interlobular sulci. Sulci of Upper Surface. The interlobular sulci (sulci intercerebellar notch.
the worm and both hemispheres into lobules; and each lobe is composed of a central and two lateral lobules. These sulci are best seen in a median section of the vermis and
lobulares} divide
are
named
worm,
(1)
viz.:
(s.
pracentralis],
which
is
located
velum
between the lingula and lobulus centralis, in the (Fig. 79). worm; between the vinculum and ala, in the hemisphere. It terminates in the horizontal sulcus. When the vinculum is
wanting the precentral sulcus is present only in the vermis. (2) The postcentral sulcus (s. postcentalis), in the worm,
separates the lobulus centralis from the culmen; and, in the hemisphere, the ala from the anterior part of the quadrangular lobule (Figs. 77 and 79). The sulcus is situated at the upper
border of the anterior cerebellar notch and runs just under the anterior border of the tentorial surface of the cerebellar hemis-
249
Both central
sulci terminate
on the dorsum
of the supe-
medullary lamina in the horizontal sulcus. (S. pradeclivis, s. primarius] (Figs. (3) Predeclivil Sulcus. and 79). Behind the culmen and anterior part of the quad76 rangular lobule, a half inch from the anterior border of the tenthere
is
tonal surface,
declive
It
the
predeclivil
sulcus.
It
bounds the
ends at the junction of the anterior and middle thirds of the antero-lateral border of the hemisphere in the horizontal sulcus.
Embryologically
it
of the cerebellum,
by Kuithan.
Its
second to appear; it is the deepest sulcus hence the name, sulcus primarius, given it development begins near the end of the third
is
month
(4)
in utero
(Cunningham).
The
79)
is
postdeclivil sulcus (s. postdeclivis) (Figs. 76 and located in the posterior cerebellar notch, from which it
curves outward and forward in the superior surface of the hemisIt separates the declivil lobe from the folium vermis, pheres.
in the
in the
hemispheres. It ends in the horizontal sulcus at the junction of the posterior and middle thirds of the antero-lateral border.
this sulcus
Being behind the crescentic gyri of the quadrangular lobule, may be called the sulcus postlunatus. It appears a
month
later
before backward:
Hemisphere.
Vinculum
Ala
Worm.
Lingula
Precentral sulcus
Hemisphere.
Vinculum
Ala
Lobulus quadrangularis,
pars anterior.
Lobulus Centralis
Posteentral sulcus
Lobulus quadrangularis,
pars anterior.
Culmen Monticuli
Predeclivil sulcus
Lobulus quadrangularis,
pars posterior
Declive Monticuli
Lobulus quadrangularis,
pars posterior
Postdeclivil sulcus
Semilunaris superior
Folium Vermis
Horizontal sulcus
Semilunaris superior
250
THE RHOMBENCEPHALON.
(Figs. 76, 77
and
79).
The
lobes
first
where the branches of the medullary body (lamina medullares) will guide the student and where the sulci are most easily identified. These lobes include the divisions
section,
median
of the
worm and of the hemispheres, and are five in number. Lingula and Vincula, Lobus Lingulae. The lingula
is
very small lobule of the vermis entirely concealed in the anterior cerebellar notch by the overhanging central lobule. It is a tongue-
gyri.
its
It
white
the lingula tapers off and is sometimes represented in the hemisphere by a very thin gyms called the vinculum lingulas. The vinculum is bounded by the
center
continuous.
Laterally,
cerebelli in front,
The
from the central lobe. Central Lobule and Alae, Lobus Centralis (Figs. 77 and The lobulus centralis is situated between the precentral and 79).
It covers the
and in turn
is
Four or
it is
five
make
up.
On
sagittal section,
seen
form a
single
vitse).
The
Culmen and Anterior Part of Quadrangular Lobules, Lobus Culminis (Figs. 76 and 79). In the culmen monticuli the
surface of the cerebellum reaches
large lobule
It is
and
made up
It is a highest elevation. occupies half of the tentorial surface of the worm. of three or four prominent gyri, which extend
its
The
rangularis occupies about one-third of the tentorial surface of the hemisphere. The predeclivil sulcus separates the culmen
culmen) from
251
bus Declivis
cerebelli.
Declive and Posterior Parts of Quadrangular Lobules, LoThe declive monticuli forms the (Figs. 76 and 79).
of the monticulus
The
Its gyri are continued into either hemisphere, where they form a large crescentic lobule, the pars posterior lobuli quadrangularis. The increased size of the lobe in the hemisphere is due to the
The
anterior
posterior parts lobulus quadrangularis, which forms the anterior two-thirds of the tentorial surface of the hemisphere. The declive and its
and
constitute the
hemispheral
extensions
are
inclosed
between
the
predeclivil
and
folii
postdeclivil sulci.
lobules, lobus
76 and
79), lies
notch.
It
common
to the postdeclivil
(Cunningham).
contains a single medullary lamina beset Rarely with rudimentary gyri, which are largely developed in the hemisit is
absent.
It
pheres.
The
is,
It
surface,
comprises the posterior third of the hemisphere's tentorial and forms one of the remarkable features of the human
cerebellum.
prominent laterally and depressed centrally (as the organ is viewed inverted), the hemispheres being separated by the anteroposterior groove, called the vallecula cerebelli (Figs. 77
and
80).
The
is
(little valley) is occupied by the inferior -worm and bounded on either side by a small cleft, 'bet ween the worm and
vallecula
The
252
inferior
THE RHOMBENCEPHALON.
cerebellar surface
is limited by the horizontal sulcus from the medulla by the transverse fissure of the separated cerebellum. It is more complex than the superior surface; and
and
is
its sulci
are
as they pass
from the
worm
Sulci of
Lower Surface
(Fig.
80).
The
interlobular sulci
79 and 80)
in the anterior
end of the
worm between
the nodule
Fig. 80.
a.
(Original.}
Pyramid, b. Flocculus, c. Nodule, d. Brachium pontis. e. Restiform body. f. Superior medullary velum, g. Brachium conjunctivum. h. Quadrangular lobule, i. Postnodular sulcus. j. Uvula, k. Tuber' vermis. 1. Ant. and m. Post. Slender lobules, n. Inferior semilunar lobule, o. Tonsil, p. Biventral lobule, q. Horizontal sulcus. r. Lobulus s. Prepyramidal sulcus. t. gracilis. Postpyramidal sulcus. u. Post, cerebellar notch.
v.
Sulcus valleculae.
In the hemisphere it winds forward and outward between the inferior medullary velum and the tonsil, and then continues lateralward between flocculus and biventral lobule to the horizontal sulcus. It is the first cerebellar sulcus to be
and uvula.
developed (Cunningham).
(2)
(s.
prapyramidalis} (Figs. 79
is
and
very con-
253
the tonsil, separating it from the biventral lobule. behind the flocculus in the postnodular sulcus.
(3)
terminates
(s.
postpyramidalis] (Figs
between the pyramid and tuber vermis, is near the It forms an oblique groove in either posterior end of the worm. sulcus valleculae, from which three concentric sulci extend into
the
79 and
hemisphere. The anterior of the three (the pregracile), usually considered the postpyramidal sulcus in the hemisphere,
separates the biventral lobule from the slender lobule (1. gracilis); the remaining two (midgracile and postgracile) subdivide the slender lobule into anterior and posterior slender, and separate
The gracilis from the inferior semilunar lobule. bounded behind by the horizontal sulcus. Sulci and lobules of the lower surface of the cerebellum, from before backward:
the
lobulus
last is
Hemisphere.
Flocculus.
Worm.
Nodule.
Postnodular sulcus.
Hemisphere.
Flocculus.
Tonsil.
Uvula.
Tonsil.
sulcus.
Prepyramidal
Biventral lobule.
Biventral lobule.
Tuber vermis.
Horizontal sulcus.
semilunar lobule.
semilunar lobule.
They
worm
hemisphere as on the upper surface (Figs. 76 and 80). Excepting in the posterior lobe, only a small ridge beneath the sulcus valleculae joins the central and lateral lobules together. The
to the inferior lobes are four in
number.
and two
the
lateral lobules as
is
composed
of a central
surface.
The
and
lobule in
worm
gives
its
name
to the lobe.
(Figs. 79
80).
The
which
worm.
It
It is
composed
medvitae.
254
THE RHOMBENCEPHALON.
larger
It
is
it is the counterpart of the lingula on the superior bounded by the sulcus valleculae on either side. The inferior medullary velum extends laterally from the nodule, and in part blends with the brachium pontis of the cerebellum.
Though
velum.
In front of the
nally to a
tonsil,
a layer of gray matter (pedunculus flocculi) That gray matter enlarges more exter-
smah
ends.
human
Embryologically, the flocculus is the oldest lobule of the cerebellum, as is the floccular sulcus (postnodular sulcus)
which bounds it, the first one formed. The flocculus is very small and rudimentary in man. It is divided into an anterior and a posterior part, the latter being called the secondary flocculus.
The
flocculus
is
lobule
by the postnodular
the nodule,
The whole
line of structures,
namely,
(Figs. 79
and
80).
The
uvula (uvula vermis) comprises a considerable part of the vermis behind the nodule. It broadens backward and is widest
Bounded on
either side
by the sulcus
vall-
eculae, projects into the valley like the uvula into the isthmus of the fauces. It comprises one large branch of the arbor vitae
its
origin into
it
hem-
From
the furrowed
band the
tonsil
expands downward
The
tonsil
rowed band, medially; and, behind, it conceals the connecting Its large size ridge between the pyramid and biventral lobule.
a prominent feature of the human cerebellum. fossa containing the tonsil is the bird's nest (nidus avis).
makes
it
The
Beis
the
prepyramidal sulcus.
(Figs.
make up
covers
2 55
one strong lamina of the arbor vitae, which divides into two near the surface. It forms the most prominent lobule of the inferior worm. A low connecting ridge joins the pyramid to the
biventral lobule in the hemisphere.
lus biventer) is triangular in outline.
The
Its
flocculus
and
is
sulcus;
its
apex
Fig. 81.
a.
(Original.)
Sup. semilunar lobule, b. Corpus medullare. c. Post, part quadrangular lobule, d. Nucleus dentatus. e. Ant. part of quadrangular lobule, f. Interior of dentate nuc. g. Central sulci. h. Brachium pontis. i. Restiform body. j. Inf. semilunar and slender lobules, k. Hilus of nuc. dent. 1. Biventral lobule.
it
composing it radiate from the apex toward the base, and are divided into two groups by a very deep
to the
pyramid.
The
gyri
intralobular sulcus.
flocculus.
nally,
Its
lateral
extension
sulcus
is
little
it
beyond the
The postpyramidal
bounds
postero-exter-
and separates it from the slender lobule. Tuber Vermis, Slender and Inferior Semilunar Lobules, Lobus Tuberis (Figs. 79 and 80). The tuber vermis forms the It resembles the lobules of posterior end of the inferior worm.
the vermis superior, because
some
of
its
half
dozen
tertiary gyri
256
THE RHOMBENCEPHALON.
are continued into the hemispheres, the sulcus valleculas not cutA bifurcated lamina of the arbor vitae enters into ting them off.
the tuber.
The
it
The
slender
and
comprise the posterior two-thirds of the inferior surface of each hemisphere, extending from the biventral lobule to the
compose the lobules. The gyri are divided into three groups by the midgracile and postgracile sulci; the anterior and middle groups are named the anterior slender and posterior slender lobules, they constitute
to fifteen gyri
postero-lateral border.
Twelve
The
posterior
is
The
of the vermis.
The
lunar lobules
bellum.
is
human
cere-
of the cerebellum
is
body
I.
(Figs. 81
and
84).
is
The cortex of the cerebellum (substantia corticalis cerebelli) made up of two thick layers visible to the naked eye, viz,, (i)
(2)
At the junc-
two layers is a single row of large pitcher-shaped cellbodies, which are characteristic of the cerebellar cortex and are often visible to the unaided eye. They are the bodies of Purkinje's cells,
and are considered in the deep part of the first macroUnder scopic layer, where they form the stratum gangliosum.
the microscope three layers are easily seen, viz., (i) the gray layer (stratum cinereum); (2) the ganglion cell layer (stratum gan-
gliosum); and (3) the granular layer (stratum granulosum). Thickest on the (i) Superficial Layer (Figs. 82 and 83).
laminae
and thinnest beneath the fissures, this layer contains small and large stellate cell-bodies with their processes, which constitute the stratum cinereum proper; and the large Purkinje cell-bodies with their dendrites and recurrent axones, together
257
ThePurkinje
cells
gangliosum.
Cells.
located
bodies of Purkinje's cells (Figs. 82 and 83) are surface of the superficial layer in the
stratum gangliosum.
longest axis.
layer,
They measure from 100/1 to 135/1 m their Each has one axone which, after piercing the deep
fiber of the
becomes a
medullary body.
It gives off, in
the
rela-
From
each cell-body antler- like processes, the dendrites, are given off; they ramify toward the surface in a wide plane at right angles to the free border of the gyrus. The edge of the plane only is seen
in a longitudinal section of the gyrus
and the arborization is very narrow and tall. The stellate cell-bodies, an outer and inner layer, together form the stratum cinereum. They increase in size
toward the Purkinje cells. They have rich dendritic processes and one axis-cylinder each. Their processes ramify throughout the stratum cinereum and stratum gangliosum.
called
The
inner
layer of the stratum cinereum contains the larger cells; they are
the
"basket cells."
Their
axis- cylinder
processes
run
parallel with the surface and at right angles to the border of the gyrus; they give off vertical branches, which descend to Purkinje's
In corpuscles and inclose them in a basket work of filaments. the outer layer of the stratum cinereum the stellate cell-bodies
are smaller than in the inner layer. They branch freely and terminate in claw-like end-tufts in contact with other stellate cells.
fibers of the superficial layer (Figs. 82 and 83) have three sources: (a) The processes of neurones within the layer,
The
stellate cells
and
the dendritic planes and recurrent collaterals of Purkinje's cells, (b) The processes of cell-bodies in the deep layer, whose T-
branched axones pierce the dendritic planes of Purkinje in the first layer; and, the processes of the large granules whose dendrites ramify
(c)
The
fibers of the
medullary
or end largely in the cellular layer. The axones of Purkinje's neurones compose all of the corticifugal fibers. They end in the cerebellar and vestibular nuclei of the cat (Clark
projection rise
17
258
THE RHOMBENCEPHALON.
The
corticipetal fibers,
which
rise
brain or in the spinal cord and ganglia, terminate in varicose fibrils chiefly in the superficial layer. These fibrils entwine about
the "primary
Fig. 82.
Section of cerebellar gyrus made parallel with its free border. Diagrammatic. (After Kolliker from Cunningham.)
G R. Small granules with claw -shaped dendrites and long axones that run out into the gray layer and divide T like. N. Axone of small granule. P. Purkinje cells seen in profile, showing border of dendritic planes in gray layer.
(2)
82 and 83)
of uniform thickness.
It
medullary projection. It contains a few superficial granules which are comparatively large in size and many small granules in which the nucleus occupies nearly the whole cell-body.
Cells of the Granular Layer. The granules are small, round, or stellate cell-bodies, largest near Purkinje's cells, closely packed externally, but scattered among the projection fibers centrally.
259
Each small granule has three to five short dendrites, which soon break up into claw-like tufts in contact with adjacent granules, and one long axone. The axone runs out into the superficial layer, branches T-like, and, piercing the dendritic planes of Pur-
Fig. 83.
Section across a cerebellar gyrus at a right angle to the free border. Diagrammatic. (Gordinier after Van Gehuchten.)
Showing large stellate cells of first layer with their basket-work endings; the cells of Purkinje, their dendritic planes in the gray layer and their axones running through the granular layer to the medullary lamina of the gyrus; two large granules of Golgi type; the small granules whose T-branches run parallel with the border of the gyrus; moss-like endings of Cajal, etc.
division.
until exhausted by multiple are dendraxones, the type of Golgi. remarkable arborizations toward the medullary
them
projection, touching
The
den-
260
THE RHOMBENCEPHALON.
(Figs. 82
and
(a)
83).
The nerve
processes of
The
the granules, (b) the axones of Purkinje's cells running down into the medullary projection, together with their recurrent collaterals,
and
most of which run through the granuend in the first layer; the remainder
terminate in the deep layer in the moss-like appendages of Cajal. The functions of the stellate cells, the "basket cells" and the
are probably receptive and associative; they receive impulses through the projection fibers and transfer those impulses to the dendrites or bodies of Purkinje's cells. Purkinje's
granule
cells
cells originate
(Gordinier), and
impulses for the coordination of muscular action for the inhibition of nervous activity in the op-
produce incoordination, chorea, athetosis and, rarely, convulsions. The neuroglia of the cerebellum is similar to that in the cere-
brum.
short-rayed cells are scattered throughout the gray while the long-rayed are located near, or within, the substance, white substance. In the region of Purkinje's cells, near the surface of the deep layer, are the bodies of the arborescent cells, whose
The
processes form a fine interlacement about the cell-bodies of PurThey kinje and then extend in parallel lines out to the surface.
form a neuroglia
basalis).
mater (lamina
II.
ganglia of the cerebellum are the nucleus dentatus, nucleus emboliformis, nucleus globosus and nucleus fastigii (Figs. 81
The
and 84). All these nuclei are made up of stellate cell-bodies, which vary in size from six to forty microns. They form relay
stations in the paths going out of the cerebellum.
(corpus dentatum)
is
imbedded
The nucleus dentatus measures lary body of each hemisphere. a half inch in length and a quarter of an inch in width (Fig. 81).
26l
with white
fibers, which issue from its open anterior and form the greater part of the brachium It also receives
conjimctivum
kinje's cells
cerebelli.
many
common
sensory
path.
are just visible to the naked eye under One of these, a club-shaped mass,
Fig. 84.
Horizontal section of cerebellum cutting nuclei and brachia conjunctiva. (Morris's Anatomy after Toldt.)
the cork-like nucleus emboliformis, partly closes the hilus of the dentate nucleus. Medial to that is an elongated antero-posterior The third ganglion, bulbous behind, called the nucleus globosus.
nucleus
is
in the anterior
end
of the
worm,
It is called
262
THE RHOMBENCEPHALON.
'
In the last two the point of the roof, nucleus fastigii (Stilling!). stellate cell-bodies are larger than in the nucleus emboliformis
or dentatus; but, otherwise, they are alike in structure. The nucleus fastigii and nucleus globosus form a part of the origin of
the acustico-cerebellar tract, which descends to the nucleus of
Deiters in the medulla; probably axones of Purkinje's cells also enter into the acustico-cerebellar tract in man, as they do in the cat (Clarke and Horsley). Perhaps the nucleus emboliformis
and nucleus dentatus have a less important connection with the same tract. The small cerebellar ganglia, and especially the nucleus fastigii and nucleus globosus, constitute a relay in the arc
of equilibrium.
contains
of the Cerebellum (Figs. 79 and 81). all the white matter of the cere-
a strong body measuring a third of an inch in thickness vertically in the middle of the hemisphere, but in the worm
is
a thin sheet and is very slender as seen in median section. branches to the cerebellar gyri are called the medullary lamina (lamina medullares}. Viewed in a sagittal section of the hemisphere, the medullary laminae are short and stubby branches of a
it
Its
very thick trunk; but the tree-like appearance of the medullary body and laminae in the vermis is perfect, hence the name, arbor
vita,
which
is
applied to
them
there.
notch the medullary body divides into a thick superior lamina and a thin inferior lamina which are separated by a transverse
furrow, the bottom of which constitutes the peak, or fastigium, of the fourth ventricle. The inferior lamina is the inferior medullary velum, already described; this, with the continuation of its
ependymal epithelium, forms the roof of the inferior half of the fourth ventricle. The superior lamina of the corpus medullare forms the three pairs of connecting bands (peduncles) and the Medullated axones make up the superior medullary velum.
entire corpus medullare
and
its
divisions.
We
II.
Commissural
fibers.
III.
Association fibers.
PROJECTION FIBERS.
I.
263
PROJECTION FIBERS.
it,
do so through
Brachium Conjunctivum
ium conjunctivum
(Figs. 44, 77
is
The
it,
brach-
(superior peduncle)
body and the brachium pontis; and, in the angle between the brachium conjunctivum and the restiform body, is the vestibular nucleus of Bechterew (the upper part of The brachium conjunctivum is joined to Deiters's nucleus).
fellow of the opposite side
its
(velum medullare superius). Two bundles of fibers run throughout the brachium conjunctivum. They are afferent and efferent.
cerebrum, which comprise nearly the whole brachium, are axones of cell-bodies situated in the nucleus dentatus. This group of fibers partially buries itself in the dorsal area of the pons, then penetrates the mid-brain and decussates ventral
First, the fibers to the
It ends largely in the the inferior quadrigeminal colliculi. opposite red nucleus, but partly in the thalamus. It constitutes
to
one segment of the indirect sensory path. In the red nucleus this path is relayed to the thalamus and to the cortical fillet, or
the relay
is
fillet.
It is
probable that a
few
fibers of this
and
fourth,
brachium terminate in the nuclei of the third and perhaps the sixth cerebral nerves. The second
fibers rises in the opposite red nucleus, pursues a reverse course and ends in the nucleus dentatus. A few fibers from
group of
chium conjunctivum and the fila lateralia pontis traverses the same part of it. Although the greater number of these fibers in the conjoined brachia decussate in the tegmentum, a few run to the red nucleus and thalamus of the same side; while the fila lateralia cross in the pons from the opposite pontine nucleus and
the
commissural
isthmus.
between Bechterew's vestibular nuclei and cross through the superior velum in the The superior medullary velum (Fig. 65) arches over
fibers
264
the
THE RHOMBENCEPHALON.
fourth
ventricle
composed
of
between the brachia conjunctiva. It is One distinct longitudinal and transverse fibers.
bundle, derived from the spinal cord, passes through it to the worm. This is the anterior ascending cerebello-spinal tract (Hoche). The decussating root-fibers of the fourth nerve (trochlear) course
transversely through
it
and
also the
commissural
fibers
between
Bechterew's nuclei.
Brachium Pontis (Middle Peduncle, Figs. 44, 78, 81 and 91). The brachium pontis comes from the pons, of which it forms the
anterior
ullary
fibers.
med-
body
brachium con-
junctivum and the restiform body. According to Klimoff fibers running to the cerebellum make up the entire brachium pontis.
These are axones of the nucleus pontis, chiefly the opposite one. Most of them run to the cortex of the cerebellar hemisphere; a small number runs to the vermis cerebelli. They form a seg-
ment
in the indirect motor path contained, above the pons, in the medial and lateral fifths and the intermediate bundle of the
basis pedunculi.
we may include
fibers.
The
latter is largely
may
glionar layer of the cortex; it terminates in the nucleus of Deiters. The principle origin of the acustico-cerebellar tract is in the opposite
nucleus
fastigii
It
descending limb of the arc of equilibrium. By far the greater number of fibers in the brachium pontis, whether efferent or afferent, are crossed fibers.
Possibly there are in the brachium pontis axones of Purkinje's cells which terminate in the nuclei pontis on both sides and in the nuclei of the reticular
formation.
78, 81
and 91) can be traced both to and from the upper part of the hemisphere and worm. Inferiorly, it is the restiform body of
the medulla oblongata.
The bundles
origin
is
of
component
fibers are
very numerous:
cerebellar tract),
(i)
The
whose
COMMISSURAL FIBERS.
and termination
(2)
265
its
in the superior
worm, forms
of
central part.
The
external
arcuate fibers
surface.
anterior)
uli
form
its free
gracilis
superior,
site
and nucleus funiculi cuneati and end in the vermis the posterior on the same and the anterior on the oppo(3)
side.
bundle from the lateral nucleus of the medulla on the same side. (4) The internal arcuof the restiform
body and, with the acustico-cerebellar tract, make up all, or the greater part, of the indirect motor path from the cerebellum to
the medulla.
According
to
Cunningham
the
cerebello-olivary
are ascending fibers. There are probably a few axones of Purkinje's cells that run down through the corpus restiforme and are continued without interruption into the anterior descending cerebello-spinal tract of the spinal cord; but surely the greater part of that tract is relayed in
the medulla and pons, chiefly in the nucleus of Deiters and in the
inferior olivary nucleus. (?)
should be noticed that the (posterior) cerebello-spinal fascicthe external arcuate fibers and the anterior ascending cerebelloulus, spinal tract, with the tract from the lateral nucleus, are the in- comIt
ing parts of the indirect sensory path, and that the brachium conjunctivum is the out-going continuation of that path up to the red nucleus and thalamus; and again, that the brachium pontis, the
acustico-cerebellar tract and the descending fibers of the restiform body constitute segments of the indirect motor path and have to do with coordination and equilibrium.
II.
COMMISSURAL FIBERS.
are joined
The
of
cerebellar hemispheres
by transverse
fibers,
which there are two sets, namely: One near the anterior end of the worm beneath the central lobe, and the other at the
posterior
ASSOCIATION FIBERS.
266
THE RHOMBENCEPHALON.
but there appears to be nothing analogous to the long association fibers found in the cerebrum.
RHOMBENCEPHALON.
SECTION
II.
THE PONS
(VAROLII).
the ventral part of the rhombenthe cerebellum being its dorsal portion. cephalon, By a transverse indentation of its roof, the posterior brain-vesicle is parti-
an upper vesicle, the metencephalon, and a lower the myelencephalon; the. latter is the embryonic medulla, vesicle, the former gives rise to the cerebellum and the pons. The pons is developed from the floor of the metencephalon (Fig. 79). It
ally divided into
is
so
named because
it
tween the mid-brain, above, and the cerebellum and medulla oblongata, below; between the medulla and cerebellum, and be-
tween the two cerebellar hemispheres (Fig. In shape the pons is roughly cylindrical.
85).
It
or ventral part, the pars basilaris pontis, and a narrower dorsal portion, the pars dorsails pontis (Fig. 87).
about one inch long. It is a little broader than long, and measures an inch, dorso-ventrally. Position. It rests in the anterior end of the groove which
Size.
is
The pons
magnum
to the
dorsum
it
sellae,
and
to the
Superiorly,
continuous
with the medulla oblongata. Surfaces of the Pons. The pons has four surfaces,
rior
viz.,
supe-
(attached); inferior
and
posterior
and two borders, namely, right and left lateral, continuous with the brachium pontis of the cerebellum. The superior and inferior surfaces are made by section, and are directly continuous with mid-brain above and the medulla
(partially free);
below.
Anterior
of the
rests
Surface
(Tuber
annulare).
The
anterior
surface
pons (Fig. 85) looks forward and slightly downward and on the sphenoid bone behind the sella turcica. It is divided
267
Fig. 85.
THE PONS
into lateral halves
artery;
(VAROLII).
269
and
is
by the sulcus basilaris, containing the basilar bounded laterally by a sagittal plane cutting the
Vertically the surface
side to side.
It
is
slightly
convex, and
is
markedly so
from
shows trans-
due
form
it
of the cerebellum.
The
end of the pons bend downward (fasciculus and form a rounded margin, which covers the lower obliquus)
Those
at the superior
part of the bases pedunculi of the mid-brain; at the inferior extremity of the pons, the fibers are convex downward and partially
conceal the pyramids of the medulla oblongata. The two roots of the fifth nerve (trigeminal) are attached to the lateral border (Henle) of this surface, a
little
The
bellum
posterior
(Fig. 88).
surface of the
It is free in its
The
ventricular area of the posterior surface is completely concealed by the superior medullary velum. If examined, it is found to be divided into lateral halves by a median longitudinal groove.
Each half presents in its posterior part a rounded eminence, the colliculus facialis, which flanks the median furrow and is in
turn bounded, laterally, by a linear valley, the sulcus limitans, lying near the brachium conjunctivum cerebelli and parallel with
it.
end of the valley is called the fovea superior; its upper part has a bluish tint, due to underlying pigmented Attached Area. Lateral cells, and is called the locus caruleus.
inferior
to this ventricular area, the posterior surface of the
The
pons
is
attached
to the restiform
of the cerebellum.
The
restiform bodies enter the surface near the lower end of the
pons and then bend downward into the medulla oblongata; while
the brachia conjunctiva, in their course up to the cerebrum, partly imbed themselves hi the lateral part of the posterior surface
issues
and form the walls of the fourth ventricle. The lateral fillet from this surface just external to the brachium conjunctivum. It runs obliquely across the upper end of the brachium to the inferior colliculus of the corpora quadrigemina, and pro-
270
duces a
flat
THE RHOMBENCEPHALON.
striated ridge,
which
may be
is
The pons
and
composed
fibers
of gray matter.
The
trans-
Fig. 86.
(Morris's
sulcus. b. Superior f ovea. c. Limiting sulcus. d. Medial eminence, e. Striae Inferior fovea. g. Nucleus funiculi cuneati. h. Taenia of fourth ventricle, k. Posterior median fissure. 1. Aquaej. Nucleus funiculi gracilis (clava). ductus cerebri. m. Nucleus incertus. n. Locus caeruleus. o. Colliculus facialis et nucleus abducentis. p. Nucleus N. cochlearis (tuberculum acusticum). q. Area acustica (nucleus vestibularis). r. Nucleus intercalatus. s. Trigonum N. hypoglossi. t. Alacinerea. u. Fua.
Median
medullares.
f.
i.
Area postrema.
niculus separans.
v.
Obex.
verse fibers are found chiefly in the basilar portion of the pons; the longitudinal, in both the basilar and the dorsal part. The
anterior longitudinal intersect the deep transverse fibers of the
pars basilaris.
271
transverse fibers form three consecutive layers in the pons, the anterior, the middle, and the posterior layer. They
area of the pons, the latter in the dorsal area (Figs. 87 and 88). The superficial transverse fibers are anterior in position
and form a thin compact layer constituting the anterior surface of the pons (Fig. 87). They are not intersected by longitudinal
fibers
;
of Pars Basilaris.
These form a
thick lamina posterior to the superficial transverse fibers and in contact with the superficial lamina. They are intermingled with
longitudinal fibers from the bases pedunculi, viz., the pyramidal, fronto-pontal, temporo-pontal and intermediate tracts (Figs.
87 and 88).
large
fibers is a
mass
of gray, matter,
on
Their origin
is
found
in the opposite nucleus pontis (Klimoff). They continue the from the termination of the fronto-pontal, indirect motor path
In connection with the superficial and deep transverse fibers pons there should be mentioned an
is
independent strand, the fila lateralia pontis, which at the upper border of the pons and buried more or
situated
less in the
isthmian furrow between the pons and mid-brain. It is called by Henle the taenia pontis. According to Sir Victor Horsley, it
from the nucleus pontis just ventral to the interpeduncular ganglion, and, winding round the isthmus, enters the cerebellum
rises
Its destination is
probably
the nucleus dentatus and nucleus fastigii (Brain, Vol. 29, No. 113). The transverse fibers of the pars dorsalis pontis (Fig. 88) compose a thin layer on the dorsum of the middle transverse
fibers,
separating
them from
the
formatio
reticularis.
This
272
transverse lamina
It is called the
is
THE RHOMBENCEPHALON.
present only in the inferior part of the pons.
corpus trapezoideum.
(corpus trapezoideum) lies in the dorsal next the boundary between the pars basilaris area of the pons, and the pars dorsalis pontis. Its fibers rise chiefly from the
trapezoid body
The
nuclei of the cochlear nerve and, after decussating in the raphe, are continued up in the lateral fillet to the inferior quadrigeminal
colliculus.
few
nerve.
The
body and
form relays
for a
number
striae,
of
its fibers.
The
corpus trapezoideum
and the
lateral
fillet,
which
is
the con-
tinuation of both, form the second stage in the auditory conduction path; and the auditory impulses are continued (a) through the
brachium quadrigeminum
inferius
and
(b) the
temporo-thalamic
The
body.
Four fasciculi make them up. (Figs. 87 and 88). The frontoare the four motor tracts of the basis pedunculi. They and intermediate tracts terminate in the pontal, temporo-pontal
nucleus pontis. The pyramidal fibers run from the middle threefifths of each basis pedunculi down through the middle transverse layer of the pons to the pyramids of the medulla oblonTogether with the above cerebro-pontal tracts they form gata. a thick bundle on either side of the median line, which presses
down
the superficial transverse fibers and produces the sulcus basilaris. The nucleus pontis, one on either side, is situated among the pyramidal fibers.
cussate
The pyramidal
which leave them to deand end in the nuclei of motor cerebral nerves. The Dorsal Longitudinal Fibers. These are contained in
273
the pars dor sails pontis in the formatio reticularis (Figs. 87 and 88). They are dorsal to the corpus trapezoideum, and lie in the floor
where they are intermingled with the ventricular gray substance. They do not form a compact layer, but are collected into a number of distinct strands, of which the
of the fourth ventricle,
naked eye in Weigert-Pal sections. The dorsal longitudinal fibers are mingled with many transverse and oblique fibers; and, thus, there is produced the netlike arrange-
reticularis.
The
the
formatio
tegmentum pons and comprises the tegmental region of the pons. The gray matter in the meshes of this network, which is the stratum nucleare continued up from the medulla, contains the nuclei of the fifth, sixth and seventh cerebral nerves and a part
of the mid-brain of the nucleus of the eighth nerve,
continued
down from
and
matio
reticularis,
viz.,
the
nucleus
lateralis
superior,
medius
and
inferior,
medius.
In the formatio
the dorsal longitudinal fibers of the pons. These tracts are as follows The anterior ascending cerebello-spinal tract, the spinothalamic tract, the medial fillet, the lateral fillet, the medial (pos:
and anterior longitudinal bundles, the rubro-spinal tract, the olivary fasciculus, and the descending root of the fifth cerebral nerve, besides unidentified fibers of the formatio reticularis
terior)
tion.
which are probably both ascending and descending in conducThe anterior ascending cerebello-spinal tract and the
descending root of the trigeminal (or fifth) nerve are the only tracts not already considered in our study of the tegmental region
of the mid-brain, page 156.
is
(lemniscus medialis, Figs. 87 and 88) a large bundle of fibers that runs through the pons next the median plane. In the lower part of the pons it lies dorsal to the
(i)
fillet
The medial
some
it.
gracilis
and nucleus
and
of
sensory cerebral nerves (Fig. 94). It conducts imIn the mid-brain it pulses of the tactile and muscular senses.
18
common
274
THE RHOMBENCEPHALON.
fillet
The medial
fillet
ends
thalamus (Fig. 47). Interruption of the medial lemniscus causes ataxia on the opposite side.
in the lateral nucleus of the
The lateral fillet (2) Lateral Fillet. (Lemniscus lateralis). forms a link in the special sense, auditory path (Fig. 89). As stated on page 160 it is but the longitudinal continuation of the
corpus trapezoideum and the medullary striae. It takes form near the middle of the pons, where the fibers of the trapezoid body bend- upward to a longitudinal direction; and it runs just lateral
Very soon it becomes separated from the medial fillet by the brachium conjunctivum of the cerebellum. It runs dorso-medially over the conjoined brachium
to the medial
fillet
(Figs.
86 and 87).
where a
end but the greater number are continued through the brachium inferius to the medial geniculate body. The chief
few of
its fibers
fillet is
though some
oideum,
lateral
It is also
of
the
fillet,
which constitute
superior olivary nucleus, and the nucleus of the partial relays in the auditory path.
fillet
from the
same
supposed to
decussate near or in the quadrigeminal bodies and terminate in the opposite inferior colliculus. Destruction of the lateral fillet
causes deafness, almost complete, in the opposite ear. (3) The spino-thalamic tract occupies the lateral part of the formatio reticularis where it is intermingled with the anterior
ascending cerebello-spinal tract (Figs. 87 and 88). As already stated, it rises in the spinal cord from the basal gray substance
of the anterior
of
common
sensory cerebral nerves in the medulla and pons. The spinothalamic tract ends in the lateral nucleus of the thalamus. It
conducts impulses of the tactile, pain and temperature senses. (4) The anterior ascending cerebello-spinal tract has the
same spinal origin and function as the spino-thalamic tract and the same course up to the isthmus rhombencephali (Figs. 87 and There, it bends backward, medial to the brachium con88).
275
junctivum, and, through the superior medullary velum, enters It thus belongs to the inthe superior worm of the cerebellum. From the cerebellar direct sensory path (through the cerebellum).
cortex the path is continued by the axones of Purkinje's cells to the nucleus dentatus, whence the brachium conjunctivum comThe anpletes it up to the opposite red nucleus and thalamus.
terior
Fig. 87.
a.
(Original.)
brachium conjunctivum. b. Formatip reticularis. cBeginning Brachium conjunctivum. d. Medial longitudinal bundle, e. Fourth ventricle, f. Superior velum, g. Descencding root of sth n. h. Spino-thalamic tract, i. Lateral medullary k. Long, fibers from basis pedunculi. 1. Superficial transverse fibers. fillet, j. Medial fillet, m. Nucleus pontis. n. Deep transverse fibers of pars basilaris pontis.
of decussation of
are the chief bundles- of a spino-encephalic system of fibers which terminates very largely in the cerebellum and thalamus, but also sends fibers to the nucleus lateralis inferior and other reticular
nuclei, to the substantia nigra, to the inferior
uli
and superior
collic-
cleus hypothalamicus,
encephalic
system
there
and the corpus striatum. In this spinoare two important paths the direct
276
THE RHOMBENCEPHALON.
tactile,
path (No. 3) and the indirect path (No. 4) for temperature impulses. It is called Cowers' s tract.
(5)
pain and
The medial
longitudinalis medians'}
(posterior) longitudinal bundle (fasciculus (Figs. 87 and 88) runs next the median
plane and just under the ventricular gray matter in a position similar to the one it occupies in the mid-brain (Figs. 47 and 48),
the pontine portion of this bundle that the fibers from the oculomotor nucleus pass to the genu of the facial nerve, ultimately to innervate the frontalis, corrugator and orsee page 156.
It is in
bicularis oculi ;
it is
and run upward through it to the oculomotor nucleus of the opposite side and make possible the conjugate movements of the eyeballs; it is also here, that fibers, which
cleus join this bundle,
the hypoglossal nucleus, leave the longitudinal bundle and enter the facial nerve at the genu to be distributed by way of the facial to the orbicularis oris. As in the mid-brain, the longitrise in
udinal bundle includes the two functional tracts, the descending strand and the ascending strand.
(6)
is
the pons in nearly the same relative position. Diverging a little from the medial longitudinal bundle as it descends through the pons, it is located
in the formatio reticularis a short distance ventro-lateral
down through
from
it.
The
normal
adult tissue only in the dorsal tegmental decussation (Meynerti) of the mid-brain: lower down it can be distinguished from the surrounding tissues by degeneration and medullation but in no
other
way
(Fig. 87).
Having given
it
fibers to the
oculomotor and
cent nerve at this level, and perhaps to other pontine nuclei (see
this tract
Rubro-spinal Tract. In the mid-brain, we have traced from the red nucleus, through the ventral tegmental
decussation (Foreli), to the opposite side, where it mingles with the lateral fillet down near the isthmus (Figs. 47 and 48). It occupies the same position in the upper half of the pons; it is close
to the posterior surface of the corpus
277
In the center of the gray part of the pons (Figs. 87 and 88). the rubro-spinal tract ends. Its funccrescent of the spinal cord
tion
is
unknown, perhaps
as
it
(fasciculus olivaris} is regarded by but the weight of evidence at present tract, many 'is in favor of a descending course. Rising in the lentiform nucleus
(8)
it
Colliculus facialis
Nucleus emboliformis
Hilus of nuc. dentatus Nuc. of becht Nuclei of sth nerve
n.
Pyramidal tract
Trapezoid body and medial
Fig. 88.
fillet
(Original.)
trapezoideum
and runs between the ventricular gray substance and the brachium conjunctivum of the cerebellum down to the middle of the pons, where it joins the principle motor root. It passes lateral to the motor nucleus situated under the locus
crescentic in section
278
caeruleus,
THE RHOMBENCEPHALON.
and runs between
it
of the ter-
minal or sensory nucleus. In the lowest part of the pons there is seen the spinal tract of the fifth nerve (tracus spinalis nervi trigemini). This is a part of its sensory root and is composed of the descending limbs of
The
spinal tract of the trigeminal nerve runs lateral to the gray sub-
between the substantia gelatinosa (Rolandi), on the inner and the vestibular root of the acustic nerve and the restiside.
PONS.
(i)
In
the interstices between the deep transverse fibers of the pars basilaris pontis and the ventral longitudinal fibers, the nuclei pontis; and (2) in the formatio reticularis, just under the fourth
ventricle
where
it
the raphe, containing the bodies of large multipolar nerve cells whose axones run through the brachium pontis of the cerebellum
to the cortex
on the opposite
is
side. It extends vertically throughout continuous with the arcuate nucleus of the medulla.
The
the basis pedunculi and the intermediate bundle of the same, and thus connects these
path.
the superior olivary nucleus, the nucleus of the corpus trapezoideum, the preolivary nucleus and the semilunar nucleus; (2) the nuclei of the formatio reticularis,
of nuclei,
viz.,
nucleus lateralis medius; and (3) the nuclei of cerebral nerves the fifth, sixth and seventh, and a part of the vestibular nucleus
of the eighth nerve.
(i)
Olivary Group.
The
279
superior) is situated in the lateral part of the formatio reticularis in the dorsal portion of the corpus trapezoideum (Fig. 88). It lies just ventral to the nucleus of the facial nerve and ventrolateral to the olivary
bundle of
fibers.
The
bodied nerve
medulla.
line.
and, in this respect, resembles the olive of the Its outline is crescentic, convex toward the median
cells;
it
In
size
it
is
microscopic.
ningham
is
The
superior olive constitutes a subordinate relay in the auditory path, receiving fibers from the cochlear nuclei of both sides and
contributing fibers to both lateral fillets (Fig. 89). The superior olivary nucleus gives off a small strand of fibers,
called the olivary pedicle,
recurrent
some
of
its fibers
end,
the remainder join the medial longitudinal bundle and run to the The pedicle forms part of an trochlear and oculomotor nuclei.
auditory-ocular reflex arc. A small accessory nucleus, called the nucleus prceolivaris, is situated just a little ventral to the superior olivary nucleus; and a
second one embraces the convexity of the nucleus olivaris superior, The latter is the nucleus semilying on the medial side of it.
lunaris.
Nucleus
of the
is
deeply imbedded in the trapezoid body ventromedial to the superior olivary nucleus (Fig. 88). Its cell-bodies
This nucleus
are scattered and, like the other nuclei of the olivary group,
it
forms a partial relay for the auditory path. This nucleus is peculiar; the fibers it receives terminate in the form of cup-shaped discs,
acustic cups,
(2)
which are
The nuclei
and
All are microscopic. They are made up of large scattered cell-bodies whose axones, dividing Tlike, are both ascending and descending in direction (Tschermak).
medius.
We may
divide these axones into two groups, a crossed uncrossed. The crossed fibers pierce the median plane and
and an become
280
THE RHOMBENCEPHALON.
longitudinal in the formatio reticularis near the ventricular gray matter and medial to the root of the abducent nerve. At that
downward.
ponto-spinal tract) pass through the substantia reticularis grisea of the medulla and the lateral area of the spinal cord throughout length they end in the center of the gray crescent in successive segments until exhausted near the end of the cord (Tschermak,
its
;
Fig. 89.
Diagram of a transverse section through the junction of the medulla and pons showing the roots and nuclei of the eighth cranial nerve and the auditory paths in the pons. (After Morris's Anatomy.)
b. Nuc. of descending root. c. Restiform 'body. d. Lat. a. Bechterew's nucleus, f. Vestibular nerve, h. cochlear nucleus, e. Ventral nucleus, g. Semicircular canals, nerve, i. Cochlea, j. Dorso-lateral nucleus (Deitersi). k. Dorse-medial nucleus. Cochlear m. Superior olivary nucleus, n. Nucleus of trapezoid body. o. Trapezoid 1. Lateral fillet,
body.
Barker). The uncrossed fibers from the reticular nuclei enter the medial longitudinal bundle of the same side and there branch
T-like.
bundle into the anterior column of the spinal cord, through which some of them continue to the end. They occupy the outer side of the anterior funiculus and end in succession in
run with
the anterior columna of gray matter (Tschermak and Barker). Just what is the destination of the ascending branches of either
J. S. Collier suggests that these tracts from the reticular nuclei should be called the
group of
fibers
crossed
tracts
281
Nerve Nuclei.
The
nervi trigemini) are two in number. The genetic or motor nucleus of the fifth nerve (n. motorius} in the pons is a continuation of the mesencephalic nucleus. It is rather close to the fourth ventricle
in the extreme lateral part of
its floor,
cseru-
as the middle of the pons, where the whole group of axones passes forward into the motor root of the nerve. Cortical Connection. The nucleus receives
It
extends as far
down
fibers from the opposite pyramidal tract and perhaps from the cerebro-pontal tracts of the same side; and sensory fibers terminate in it from the sensory root of the fifth nerve, and from the terminal nuclei of other common sensory nerves, through
motor
the medial longitudinal bundle and establish its reflex connections. The terminal or sensory nucleus (n. terminalis or sensibilis)
of the trigeminal nerve begins at the middle of the
pons and
extends to the second segment of the spinal cord. At its superior end it is ventro-lateral to the motor nucleus and under cover of
the brachium conjunctivum of the cerebellum (Figs. 86 and 87). Near the medulla it lies ventro-medial to the restiform body and the vestibular root of the acustic nerve.
This part of
it
is
almost
and
its
distance
from the
is
ventricle
is
greater than
gelatinous in character
posterior
in the
is
cord.
It
receives
the
whole sensory root of the trigeminal nerve. Just lateral to it runs the spinal tract of the fifth nerve, the fibers of which gradually
bend into the nucleus and terminate from the nucleus pursue several
fibers
in rich arborizations.
different
Axones
Reflex
courses:
(a)
go directly to the
motor nucleus
of the fifth
fibers are supposed to enter the opposite through which they reach the thalamus;
perhaps, some run through the anterior cerebello-spinal tract cerebellum, (c) Tactile and muscular sense fibers proceed to the thalamus, probably through the medial fillet on the
opposite side.
Nucleus
of the
This
282
THE RHOMBENCEPHALON,
is
motor nucleus
the
close to the
ependyma
The
run lateral
on
its
dorsal surface
and then
nucleus
return lateral to
it.
Cortical Connection.
The abducent
motor
fibers
and from the cerebro-pontal tracts. It receives reflex impulses through the anterior and medial longitudinal bundles and the pedicle of the superior olivary nucleus and perhaps, also, through the brachium conjunctivum from the cerebellum. The axones of the cell-bodies in the adbucent nucleus run in two directions: The greater number run ventro-lateralward and emerge
;
at the
on the opposite
side
by way
;
The former
inner-
on the same
side as the
nucleus the latter, through the third nerve, innervates the internal rectus of the opposite eye, though that muscle receives independent
fibers
from the
third, also.
of
The nucleus
is
It is situated deep in the pons, in genetic or motor (Fig. 88). the lateral part of the formatio reticularis, beneath the superior fovea. Immediately ventral to it is the superior olivary nucleus,
and the substantia gelatinosa (Rolandi) lies dorso-lateral to it. The nucleus is placed midway between the spinal tract of the fifth nerve and the Cortical Connections. olivary fasciculus. It receives voluntary motor impulses from the cerebral cortex
of the opposite hemisphere via the pyramidal tract; and, probably,
fibers of
its
establish
motor connections.
the cerebro-pontal tracts terminate in it. These The reflex, connections of the
from the spinal tract of the trigeminal nerve, from the trapezoid body (Cunningham), and from the medial longitudinal bundle. The axones of the cellfacial nucleus are established
by
fibers
all
divided into three parts, viz., two By distinct parallel parts, joined by a very short ascending portion.
direction this root
is
283
The
recurrent part, the pars prima, runs dorso-medianward and then dorsal to the
lower end of the abducent nucleus; (2) it then ascends about onefifth of an inch (Cunningham) between the ventricular ependyma,
dorsally,
and the abducent nucleus and medial longitudinal bundle, ventrally, and this part is called the genu internum; and (3)
the pars secunda, bending sharply outward over the nucleus of the sixth nerve, then plunges ventrally through the pons this emer;
gent part of the root runs between the nuclei of the facial and trigeminal nerves. The root of the facial nerve is joined at the
genu internum by fibers from the medial longitudinal bundle which rise in the oculumotor and hypoglossal nuclei and supply
the facial muscles above the orbit
pectively.
and
is
the.
which
Its
is
said
to
to
axones
run
its
medius and
They are secretory and vasoand sublingual glands. Auditory Nerve (N. Nervi Vestibis
This nucleus
made up
The
root,
descending
and
(3) the
in position.
The
superior parts of Schwalbe's and of Deiters's nuclei extend into the pons just medial to the restiform body, and the nucleus of Deiters is prolonged dorsally along that body toward the cere-
bellum.
is
called
shall recur to the vestibFlechsig's or Bechterew's nucleus. ular nucleus in the medulla where the greater part of it is located. Lesions in the pons are usually attended by crossed paralysis.
We
The
paralysis and anaesthesia of parts supplied by spinal and by bulbar cerebral nerves are on the opposite side, but the fifth, sixth and seventh cerebral nerves of the same side as the lesion are apt to be involved. If the spino-thalamic and anterior ascending cerebello-spinal tracts are involved and not the medial fillet, the pain and temperature sense is lost, but there is no ataxia; if
284
the medial
anterior
fillet
THE RHOMBENCEPHALON.
be destroyed and not the spino-thalamic and
cerebello-spinal
tracts,
ascending
temperature sense is intact, but the muscular sense is lost on the opposite side of the body. The tactile sense is impaired in both
cases.
A
side
lesion
of the
trapezoid
fillet,
same
when the nucleus of the sixth nerve is affected; and strabismus when the root fibers, but not the nucleus, are involved. The strabismus is external if the lesion
Conjugate deviation occurs
be
irritative
and
internal
if
Des-
paralysis of the face, the frontalis, corrugator, orbicularis oculi and orbicularis oris not being affected. Complete facial paralysis
occurs
if
genu internum.
RHOMBENCEPHALON.
SECTION
III.
(MYELENCEPHALON.)
Situation.
is
It may be regarded as the part of the brain (Figs. 21 and 27). intra-cranial portion of the spinal cord, hence the expanded synonym, spinal bulb. It occupies the basilar groove of the
and
is
foramen magnum.
concealed in the valley of the cerebellum. wind forward around it, and form the basilar at
the pons.
Size.
is
junction with
The medulla
is
a half-inch thick.
Its
about an inch long, and dorso-ventrally, width at the lower end is a half-inch, also.
At the upper extremity it measures from three-quarters of an inch to one inch in width (Figs. 85 and 86).
Its
of a cone flattened
which
is
truncated apex of the frustum, nearly circular in outline, is continuous with the spinal
The
285
On the anterior cord; and the flattened base joins the pons. marks the boundary between the surface, a transverse groove
medulla and pons.
is a bilateral organ composed and 86). In the interior, the two halves are united by both gray and white matter in the raphe, but on the surface they are partially separated by the anterior and the posterior median fissures (fissura mediana anterior and f. m. posterior). These fissures are continued through the spinal
The medulla
The
anterior
median
forming the
Fig. 90.
Section of embryonic medulla. Length of back, 9.1 (Gordinier and Minot after His.)
lip.
mm.
RL. Rhomboid
X. Vagus nerve.
decussation of the pyramids; while only through the lower half of the medulla does the posterior median fissure extend.
Origin.
is
embryo (Figs. 90 and 118). The myelenand walls thicken and form the greater part of the medulla. Inferiorly, the roof undergoes some thickening; but it stretches out into a single layer of epithelium, superiorly, which is continuous at its upper end with the inferior medullary velum
encephalon
the
cephalic floor
of the cerebellum.
Ventricle.
persists in the
The common
mature
cavity of the posterior brain vesicle brain as the fourth ventricle (Figs. 118,
is,
90 and 86).
The
fourth
(see
rhombencephalon
page 313).
286
THE RHOMBENCEPHALON.
SURFACES.
The medulla oblongata
posterior
and
presents four surfaces: The anterior, two lateral, separated by the anterior lateral and
In the upper medulla, the surfaces are less distinct as they descend to
the inferior
and nearly
and
86).
The anterior
its
lateral sulcus
No
corresponding
the anterior lateral groove issue groove the roots of the hypoglossal nerve and the anterior root of the first cervical nerve. The abducent (or sixth) nerve rises nearly
in line with
it
From
The posthe lateral from the medulla separates It descends between the olive posterior surface (Figs. 86 and 87). and the restiform body and is continued through the spinal cord.
Posterior Lateral Sulcus.
terior lateral sulcus of the
cord run the posterior roots of the the sensory roots of the vagus and spinal nerves, and, likewise, glossopharyngeal nerves run through it into the medulla; while
Through
root of
the eleventh nerve emerge from the medulla through the posterior lateral sulcus. The roots of the seventh, eighth and intermediate nerves are found at the superior end of the sulcus in the
The posterior transverse groove between the medulla and pons. lateral sulcus is not parallel with the axis of the medulla, but bends
outward and forward as
for a short distance
it
ascends.
Inferiorly,
it
is
obliterated
by the crossing of the fasciculus cerebellospinalis (direct cerebellar tract) from the lateral to the posterior
surface.
by
(fades anterior} of the medulla, bounded the anterior lateral sulcus, extends from the trans-
verse sulcus below the pons down to the spinal cord (Fig. 85). It is made up of symmetrical halves united, below, by the decussation of the lateral (crossed) pyramidal tracts; but separated, above, by the anterior median fissure, which terminates at the
SURFACES.
inferior
287
Vicq d'Azyr).
On
either
side of the
median
rior surface presents a fusiform eminence, most prominent near the pons, called the pyramid. The pyramidal tract, which we
have already traced through the internal capsule, basis pedunculi and pons, forms tEe pyramid of the medulla. In the lower part
of the
(or crossed)
and
the former comprising about the medial four-fifths and the anterior pyramidal tract the lateral one-fifth of the pyramid.
dal
tract,
Transverse
fibers,
seen crossing the pyramid from within outward. They form a more or less continuous sheet of fibers, which emerges from the
anterior
median
fissure
to the
posterior 'surface,
where
The
anterior surface
is
two anterior
There bounded by and is inclosed between the roots of the hypoglossal nerve, ventrally, and those of the ninth, tenth and the cerebral portion of the eleventh, dorLateral Surface. (Fades
lateralis,
Figs. 81
and
85).
Each are two lateral surfaces, a right and a left. the anterior lateral and the posterior lateral sulcus;
is
sally.
Lateral surface
eral area.
by the
lateral
synonymous with the surface of the latis formed, above, by the olive, below, column and, winding backward over both, are the
is is
The
by the
olive (oliva)
an elongated eminence, a
(Fig. 85).
half
inch in
It is
and, superficially, is composed of fibers continuous with the fasciculus proprius of the lateral column in the spinal cord (Fig. 93).
(Funiculus lateralis, Fig. 85). It is made of three great bundles of fibers (Figs. 93, 94 and 95): The lateral fasciculus proprius, which, splitting into a superficial and
Lateral Column.
up
288
thalamic
THE RHOMBENCEPHALON.
At tract, which runs up the posterior lateral groove. the junction of the medulla with the spinal cord the cerebellospinal fasciculus (direct cerebellar tract) passes from the lateral to the posterior surface. The anterior external arcuate fibers,
running from the anterior surface backward to the restiform body, may be so numerous as to conceal the lateral column and lower
part of the olive.
(fades posterior] of the medulla comthe surface inclosed between the diverging posterior
(Fig. 86).
It
lateral sulci
Inferiorly,
median
(Figs.
fissure,
91,
is divided into lateral halves by the posterior and presents four bundles of fibers in each half 94 and 96). From the fissure outward they are as
it
follows:
The
funiculus
gracilis,
funiculus
cuneatus,
tractus
spinalis n. trigemini,
and the
is
cerebello-spinal fasciculus.
The
fun-
the posterior column of the spinal cord, and the funiculus cuneatus is in direct continuity with the lateral tract in the same column
of the cord.
The
The
fasciculus cerebello-
column
to the cerebellum
Superiorly, the posterior surface, on either side, is formed by a large rounded band of fibers, the restiform body (Figs. 91 and 96).
(corpus resli forme) is continued upward to the pons and then bends backward into the corpus medullare of the cerebellum in connection with which it has already been
studied.
Of
spinal fasciculus,
fibers are
the superficial fibers in the medulla, the cerebelloand the anterior and posterior external arcuate
we have
already learned, the restiform body contains, in addition to the above three tracts, the tract from the lateral nucleus and the cerebello-olivary
tract.
single
SURFACES.
stretches
289
That
is
continuous with the surface layer of the ependyma lining the fourth ventricle. By it the posterior surface is completed.
(Figs.
of triangular shape;
directed
medullary velum of the cerebellum; its apex, which downward, terminates at the obex and covers the
Frenulum
veli
Fourth nerve
LateraL'fillet
Superior medullary
velum
Brachium
conjunct! vum
Brachium
pontis
Restiform body
Taenia Epithelial roof of fourth ventricle
velum
Inferior medullary
Cuneate
tubercle'
Chorioid plexus
Median aperture
(Magendi)
Obex
Fig. 91.
Roof and
and
its
chorioid plexuses.
it is
attached
The
of
obliquely
then, transversely outward, inferior borders of the epithelial lamina become thickened by .the addition of neuroglia, and are in continuity with the ependyma of the ventricle. The thickened apex of the
to the lateral recess.
The
epithelial
19
lamina
is
investing
2QO
it,
THE RHOMBENCEPHALON.
tela of the fourth ventricle, this roof epithe-
median line near the obex by a foramen, the median aperture (apertura mediana ventriculi quarti, Magendi) and over each lateral recess by the lateral aperture (apertura later alls ventriculi quarti, Key and Retzii). These foramina establish communication between the subarachnoid space and the ventricle. On either side of the median line there is a longitudinal invagination of the epithelial lamina into the ventricle and a similar transverse one just below the inferior medullary velum, both of which are occupied by a vascular fold of pia mater. This
lium
perforated in the
fold constitutes the chorioid plexus of the fourth ventricle (plexus
chorioideus ventriculi
away, as
it
usually
is
If the roof epithelium be torn quarti). with the pia, a rough line of separation is
That
Two
layers of
ependyma form
the roof epithelium is thus removed, the lower triangle of the floor of the fourth ventricle is brought into view (Figs.
When
86 and 96). Notice the median longitudinal furrow bounded by the eminenticB mediates, which form the calamus scriptorius; then
the
little
gonum
and,
whose superior angle is the large lateral area, located above the ala cinerea external to the eminentia medialis, and crossed by the nearly
vagi),
last,
strice.
transverse medullary
tica.
This region
is
made up
of white
to-
(Figs.
bound ventrally and laterally the inferior part of the fourth and surround the upper extremity of the sixth ventricle and 95). 92
of the
medulla
is
continu-.
ous with the longitudinal fibers of the pons and restiform bodies, above, and with the spinal cord, below; the bulbar roots of the
eighth to the twelfth cerebral nerves, and
many
decussating or
2QI
Substantia Reticularis.
matter
is
deep parts of the medulla, the white matter enters into a great network called the substantia reticularis, which has gray matter in its meshes (Figs. 92 and 94).
above, with the reticular formation of the pons, with the fasciculi proprii of the spinal cord. The and, below,
It is continuous,
in the
many
intrinsic neurones,
for the
most
part,
which form a frequently interrupted and, a crossed ascending and descending tract.
retic-
They are chiefly the arcuate -fibers. The distinct tracts of longitudinal fibers contained in it will be noticed later
(page 293).
is
The gray
composed
and the
olivary nuclei.
Raphe (Figs. 92 and 94). The raphe is, primarily, a sagittal lamina of neuroglia derived from the floor-plate of the myelencephalon. It lies in the median plane and joins the lateral halves of the medulla together. It is very distinct hi the superior part of the medulla. Below the level of the olive, it is entirely obliterated by the
fillet
The raphe
is
pierced transversely by decussating and commissural fibers and is traversed dorso-ventralward by the anterior external arcuate
fibers.
of the
medulla
is
Transverse
fibers;
and
The transverse
fibers of the medulla are really more or and most of them are arched. They in-
clude the fibers of the following: (a) The pyramidal decussation (decussatio pyramidum), (b) the medial fillet in the decussatio
lemniscorum,
(c)
on the
is
sur-
and
The pyramidal
THE RHOMBENCEPHALON.
cated in the lower half of the medulla.
it,
About four-fifths of the and becomes the lateral pryamlower level of the pyramidal
The
decussation marks the boundary between medulla and cord. The medial fillet, the anterior and the posterior external arcuate fibers all rise in the nucleus funiculi gracilis and nucleus funiculi cuneati.
fillet
bends
inferior olives,
hence
its
name
anterior ex-
continuing their ventral direction, issue from the anterior median fissure and anterior surface of the
fibers,
arcuate
medulla
(Fig. 93),
the restiform body, through which they enter the cerebellum. The posterior external arcuate fibers run through the restiform body to the cerebellum without decussating. The external arcuate fibers, like the medial
fillet,
and muscular
(Fig. 92).
It
senses.
The
medulla.
and perhaps
in the
cerebellar ganglia;
nucleus.
terminates in the opposite inferior olivary This tract probably belongs to the indirect motor path,
five
though Cunningham believes it to be ascending in direction. (2) The dorso-ventral fibers of the medulla are found in
median raphe, (b) in either half dulla between the anterior and lateral areas, running
situations: (a) In the
series of ten or twelve fascicles
of the
me-
in a linear
lateral sulcus,
and
(c) in several
series,
which run
inward or outward through the posterior lateral sulcus between the lateral and posterior areas of each side.
(a)
The
nal arcuate
dorso-ventral fibers of the raphe are the anterior exterThese can be traced to the cortex fibers (Fig. 92).
Their origin
(Fig.
is
in
the
nuclei
funiculi
gracilis
and
funiculi cuneati
94).
Some of them seem to be interrupted in the arcuate nucleus on the anterior surface of the pyramid.
293
The
constitute
root-bundles of the eighth to the twelfth cerebral nerves the remaining groups of dorso-ventral fibers. By
is
The
from
the ventricular gray matter, near the median line, ventro-lateralward to the anterior lateral sulcus, where they emerge. Inclosing
between them and the raphe, the anterior area, they also separate The anterior and lateral areas are bounded it from the lateral area.
dorsally by the thick sheet of gray matter in the floor of the fourth ventricle.
(c)
The
glosso-pharyngeal and vagus and the cerebral root of the accessory nerve form the third group of dorso-ventral fibers (Fig. 93).
The motor fibers of the ninth, tenth and cerebral part of the eleventh nerves take their origin in nuclei of the medulla and emerge from the posterior lateral sulcus; while the sensory fibers of the vagus, glossopharyngeal and vestibular nerves enter that
sulcus
nuclei in the ventricular gray matter. and terminal, are located lateral to
from without and run through the medulla to their terminal These nuclei, both genetic
roots rising or terminating in
The nerve
from the
The posterior area comprises everything posterior area. dorsal to the above roots of the eighth to eleventh cerebral nerves. It thus includes the gray matter in the floor of the fourth ventricle,
called the stratum nucleare.
(3)
Longitudinal Fibers.
In the medulla, the longitudinal continuations of the same in the pons and the
restiform bodies; they are also continuous with the tracts of the spinal cord. They can be best located by reference to the three
areas
lateral
bounded by the above dorso-ventral fibers, namely, anterior, and posterior areas, which are distinctly outlined in the upper half of the medulla (Figs. 92 and 93). Longitudinal Fibers of the Anterior Area. The anterior area of the medulla lies betwen the raphe and the roots of the hypoglossal nerve, and between the anterior surface and the gray matter in the floor of the fourth ventricle (Figs. 92 and 93). It contains the pyramid, the medial fillet, the medial and anterior
294
longitudinal
THE RHOMBENCEPHALON.
bundles,
the
substantia reticularis alba and two and the medial accessory olivary nucleus. Excepting that part forming the lateral pyramidal tract and the medial fillet, the anterior area is continued in the anterior column of the spinal cord. It is naturally divided into a ventral and a
nuclei, the arcuate nucleus
of the
(pyramis) with the arcuate nucleus imbedded in fibers winding over it, occupies the ventral portion of the anterior area (Figs. 92 and 94). It is the continuation of the pyramidal tract and is composed of the axones of
it
The pyramid
cerebrum.
The
pyramidal
about the
of
its
In the lower part of the medulla, the pyramid breaks up into the anterior (direct) pyramidal tract (10 per cent, of the pyramid), which descends along
nerve nuclei.
the anterior median fissure in the
same
and the
pyramidal tract (80 per cent, of the pyramid), which, after decussating with its fellow through the anterior median fissure, runs down in the lateral column of the opposite
lateral (crossed)
side of the cord (Fig. 95). The fibers of the anterior tract cross in succession to the opposite side of the cord, through the white anterior
pyram-
tract for
the gray matter. Thus the pyramid forms a crossed cerebral motor cerebral and spinal nerves. small number of
lateral ward
from the
weakness
lateral funiculus of
They account
for the
on the well side, and for slight motion on the paralyzed side, which are commonly observed in hemiplegia. If, as E. A. Schafer
claims, the fibers of the pyramidal tracts end in the posterior columna of gray matter, then at least one neurone intervenes
between them and the motor neurones of the spinal nerves; but
without doubt, they enter into either direct or indirect relations with those neurones.
295
The
of very
tia reticularis
dorsal part of the anterior area is occupied by the substanalba (Figs. 92 and 93)* It contains the bodies
few nerve
cells
and
is
of a light color.
The
medial acces-
sory olivary nucleus is imbedded in it near the pyramid and among the fibers of the fillet; and three definite bundles of longitudinal
Fig. 92.
a.
(Original.")
Hypoglossal nucleus, b. Vestibular nucleus, c. Tractus solitarius. d. Restiform body. e. Substantia reticularis. f Hilus of olivary nucleus containing cerebello-olivary fibers, g. Anterior lateral sulcus. h. Pyramid, i. Anterior median fissure, j. Anterior longitudinal bundle, k. Medial longitudinal bundle. 1. Nuc. tractus spinalis n. trigemini. m. Tractus spinalis n. trigemini. n. Lateral cochlear nucleus, o. Ventral cochlear nucleus. p. Ascending anterior cerebello-spinal, spino-thalamic, and rubro-spinal tracts, q. Posterior lateral sulcus. r. Medial fillet, interolivary stratum, s. Anterior external arcuate fibers. t. Arcuate nucleus.
.
fibers
it,
fillet
and the medial and anterior longitudinal bundles. Medial Fillet (Lemniscus Medialis}. Just dorsal to the pyramid
in the anterior area of the medulla at the level of the olives, is a
fillet
296
(Figs. 92
it is
THE RHOMBENCEPHALON.
and
94).
medial
gracilis
The medial
rises
funiculi
and nucleus
through the median raphe in the fillet decussation. As the fillet runs brain-ward, it receives fibers from the terminal nuclei of
common
Along
in the
its lateral
sensory cerebral nerves and from the vestibular nuclei. border, it is accompanied for a short distance
lateral
fillet.
pons by the
The medial
fillet
is
composed
of ascending axones
sensory fibers of spinal and cerebral nerves. It carries ordinary sensations (tactile and muscular) to the superior quadrigeminal colliculus by the superior fillet, and to the thalamus by the greater
part of the medial
fillet.
The
medial
longitudinal
bundle
(fasciculus
longiiudinalis
in the mid-
mediates) (Figs.
brain and pons, constitutes a very distinct strand in the superior half of the medulla but below the level of the olive it can be identi;
degeneration. anterior fasciculus proprius of the spinal cord. Its location is next the median raphe and the ventricular gray substance, immediately anterior to the hypoglossal nucleus, in the
medullation or of
its
It
The same
It is
position occupied by here in the medulla that the hypoglossal fibers are supposed to enter it and run up to the colliculus facialis, where they join
it
is
in the mid-brain
At the middle
of the
medulla
fillet
pushes
this
what away from the median plane, so that it runs between the fillet decussation and the medial Below accessory olivary nucleus.
the level of the
fillet
decussation
it
pyramidum and
substance.
the isolated head of the anterior columna of gray Rising primarily in the gray matter of the cord, the ascending part of the medial longitudinal bundle is augmented in the medulla and pons by fibers from the terminal nuclei of sen-
sory cerebral nerves. Most of its ascending fibers cross the median line and terminate in the motor cerebral nuclei on the
297
opposite side; these form the middle links in many reflex arcs; a few decussate in the posterior commissure and end in the thala-
mus.
The
conduction
fibers.
The
descending
Fig. 93.
a.
(Original.)
Nucleus of iath n. b. Vestibular nucleus, c. Tractus solitarius. d. Nucleus ambiguus. e. Tractus spinalis n. trigemini. f. Medial accessory olivary nuclei, g. Posterior lateral sulcus. h. Ant. external arcuate fibers, i. Fasciculus proprius. j. Arcuate nucleus. k. Anterior longitudinal bundle in substantia reticularis alba. 1. Medial longitudinal bundle. m. Nucleus alae cinerea. n. Taenia of 4th ventricle, o. Restiform body. p. Gelatinous substance, q. Substantia reticularis grisea nucleus lateralis inferior, r. Ascending ant. cerebello-spinaj, spino-thalamic, and rubro-spinal tracts, s. Dorsal accessory olivary nucleus, t. Inferior olivary nucleus, u* Pyramid, v. Medial fillet, interolivary stratum.
part of the medial longitudinal bundle is the medial ponto-spinal tract of Collier (see pages 280, 360 and 373). The anterior longitudinal bundle (Figs. 92 and 95) descends
as a distinct strand,
colliculus of the
298
THE RHOMBENCEPHALON.
corpora quadrigemina, to the lower part of the medulla; there it approaches the medial longitudinal bundle and is continued along
the fissural surface of the anterior column in the cord.
Its ter-
mination
centers.
in the central gray substance, chiefly the cilio-spinal It forms the middle link in the visual reflex arc. Its
is
pages 158, 276 and 355). Longitudinal Fibers of the Lateral Area. The contents of the lateral area (Figs. 92 and 93) are as follows: Superficially, the lateral fasciculus proprius, the anterior descending, and the anterior ascending cerebello-spinal tracts and the spino-thalamic tract. Deeply, lies the substantia reticularis grisea. Imbedded
inferior
fasciculus
and the accessory olivary nuclei, and in the proprius is the main inferior olivary nucleus. The
gray matter of the substantia reticularis grisea is a part of the disintegrated anterior columna of the cord and, unlike that of
the anterior area,
cells.
it
many
large nerve
Lateral Fasciculus Proprius (Fasciculus Lateralis Proprius, Figs. 93 and 94). The whole lateral column of the spinal
cord, except the lateral pyramidal and cerebello-spinal tracts, is continued into the lateral area of the medulla. Composed of as-
segments of the spinal cord, the lateral fasciculus proprius ascends into the lateral area of the medulla, and runs in part beneath and in part superficial to the inferior olivary
ciative for different
nucleus; beyond the olive, it is continued in the substantia reticularis grisea of the medulla and reticular formation of pons and
mid-brain.
Among
the fibers of the lateral fasciculus proprius, is the triangular tract of Helwig and, dorsal
The former
is
believed to rise in
it.
They
are
made up
form
299
it
comprises
the acustico-cerebellar tract and, possibly, the cerebello-olivary tract; and, therefore, is almost completely relayed in the vestibular nuclei, the nuclei of the reticular formation, and, possibly,
94 and 95). It forms part brachium pontis and restiform body through which it reaches the medulla. In the lateral area, it descends along the anterior
in the inferior olivary nucleus (Figs.
of the
columna
corresponding area of the cord. It ends of gray matter. It was once regarded
the cord, medulla and pons. Near the isthmus the former turns backward through the superior medullary velum and ends in the
cortex of the superior vermis cerebelli; while the spino-thalamic
tract continues in the original direction, to the lateral nucleus
of
the
thalamus.
The common
up
of these
anterior
on the opposite
the
white anterior commissure, it ascends, mingled somewhat with the fibers of the above descending cerebello-spinal tract, along the lateral surface of the cord (Barker). It runs beneath
the posterior lateral groove of the medulla and through the formatio reticularis of the pons, to the point of division near the isthmus rhombencephali, whence the two divisions proceed
and thalamic terminations, as above stated. The tract is probably reinforced in the medulla and pons by the addition of axones from the opposite terminal nuclei of common
to their cerebellar
sensory cerebral nerves; and, rising primarily in relation with the posterior roots of spinal nerves, it thus forms a crossed path for common sensations, spinal and cerebral. It conducts tactile,
300
THE RHOMBENCEPHALON.
tract
is
The rubro-spinal
rior olive in the
and
it
95).
Running dorsal
to the infe-
mingles with the fibers of the anterior tract. It is continued down the cord, ascending cerebello-spinal
in the lateral
medulla
lumbar segment
(see
pages 162,
The longitudi-
nal fibers of the posterior area form many bundles and the bundles are different in upper and lower medulla. The substantia reticularis
is
small. The ventricular gray substance belongs to this area. The lower medulla contains: The funiculus gracilis, funic-
ulus
cuneatus,
tractus
spinal tract,
named from
(Figs. 94 and 96). body and the spinal tract of the fifth cerebral nerve at the surface and the tractus solitarius in the interior (Figs. 92, 93 and 96). The funiculus gracilis is the superior end of the ascending postero-medial column (GolPs column) of the spinal cord. Near its extremity it expands and forms the clava, and then tapers off and disappears along the side of the fourth ventricle. The clava is due to the nucleus funiculi gracilis, in which the fibers of the column end. The funiculus gracilis is composed of ascending branches of the posterior roots of the spinal nerves, which enter the cord below the seventh or eighth thoracic segment. Funiculus Cuneatus (Figs. 94 and 96). It is separated from
;
nervi trigemini and cerebellothe posterior median fissure outward In the upper medulla are: The restiform
spinalis
the posterior median fissure by the gracile bundle; and is the continuation of the ascending postero-lateral column (Burdach's
column) of the spinal cord. It ends about the cells of the nucleus funiculi cuneati and accessory nucleus funiculi cuneati, which
The
fibers of the
funiculus cuneatus are ascending branches of the posterior roots of the spinal nerves. The nerves contributing to this column
The funiculi are the cervical and the six or eight upper thoracic. gracilis and cuneatus carry to the nuclei of these columns common
sensations belonging to the tactile
and muscular
senses.
Inter-
301
Spinal Tract of theTrigeminal Nerve. (Tractus spinalis nervi It forms a narrow strip of the postrigemini, Figs. 92 and 96).
terior surface of the medulla,
which
is
body and tapers downward toward the spinal cord. It is composed of the descending fibers from the sensory root of the trigeminal nerve: the sensory fibers of this nerve on entering the pons divide
an ascending and a descending branch, and the debranches form the spinal tract of the nerve, which for a scending short distance is visible on the surface of the medulla. This
T-like, into
tract is continued
The
nucleus of the spinal tract of the trigeminal nerve, over which it runs and in which it terminates, is but the continuation of the gelatinous substance of the posterior columna of gray matter in the cord in the upper medulla it is situated ventro-medial to the restiform
;
body.
The
The cerebello-spinal
94 and 96) in the lower medulla crosses the posterior lateral groove and the spinal tract of the fifth nerve, going from the lateral column of the cord to the posterior area of the medulla; it then
ascends to form a considerable part of the restiform body. It takes its origin from the nucleus dorsalis (Clarki) in the spinal
cord. of
It ends,
the
superior
worm.
The
cerebello-spinal
tract
pulses received
from
viscera.
under the muscular sense, as they seem to concern equilibrium and orientation. They are sympathetic sensory.
(Corpus restiforme). In the upper medulla, forming the lateral part of each posterior area, is a large rounded bundle of fibers, called the restiform body (Figs. 92, 93 and 96).
It is the largest
Restiform Body.
it
to the cerebeltracts
(i) the
lum.
It is
made up
and
Ascending
cerebello-spinal tract;
anterior
and
posterior;
from the
Descending
tracts
and
(5)
302
je's
cells
THE RHOMBENCEPHALON.
which enter into the descending cerebellar
page 264).
tract
(?)
(see cerebellum,
The
restiform
body
is
and
coch-
nerve (eighth) (Fig. 89). Ventral to it and between the roots, is the ventral part of the cochlear nucleus ; on its lateral surface and among the fibers of the lateral
lear roots of the auditory
root, is the dorsal, or lateral part of the
same nucleus.
The ves-
the chief dorso-medial nucleus (Schwalbe), the dorsolateral (Deiters), the superior (Flechsig and Bechterew) and the nucleus of the descending root are situated dorsal and medial to the restiform body.
tibular nuclei
Tractus Solitarius (Figs. 92 and 94). The solitary tract is a small, round bundle imbedded in the middle of the ventricular
In Weigert-Pal sections of the upper medulla it is clearly visible to the naked eye. It is formed by the sensory roots of the nervus intermedius and the
gray matter of the posterior area.
ninth and tenth cerebral nerves.
It
and
in the cord.
along the medial aspect of the posterior columna of gray matter It ends on both sides of the median line in gelat-
inous gray matter, called the nucleus of the solitary tract, which surrounds it. The nuclei of these tracts converge downward;
according to Cajal, they are united near their inferior extremities by a central mass, called the nucleus commissuralis (Barker).
The
dulla
solitary tract
may be
down
hence the
and that
of the
downward from
the pons;
and
(3) of the special nuclei of the medulla, which are not represented in the pons or in the spinal cord. (i) Nucleus of the External Arcuate Fibers (Figs. 92 and
303
This
is
It
mass on the ventral and medial surface of the pyramid. Running over and through it there are the anterior external arcuate The arcuate fibers, for certain of which it constitutes a relay.
nucleus
is
It is
brought
pons by
the disappear-
fibers.
The gray substance in the substantia reticularis and in the immediate floor of the fourth ventricle (Figs. 92 and 93)
is
and
continuous with the same in the pons and mid-brain, above; is represented in the spinal cord by the H-shaped column
of gray matter. By the posterior and lateral expansion of the neural canal, in the upper half of the medulla and in the pons,
columnae of the
H-shaped column of gray matter are pushed outward to a transverse direction; and the entire bases of the anterior columnas are
brought into the floor of the ventricle. The expansion of the canal, together with the decussation of the lateral pyramidal tracts through
the anterior columnae and of the medial
terior
fillets
follows
Anterior Columna.
From
is
derived a column of
the hypo glossal nucleus (Figs. 92 and 93), two-thirds of an inch in length and extends along the
cells,
It is
upper medulla, beneath the eminentia mediacontinued into the lower medulla as far as the pyramin the
idal decussation.
By commissural
fibers
it is
gives origin to the hypoglossal nerve proper; and probably to a small fasciculus which by way of the medial longitudinal bundle
and supplies the orbicularis oris. The hypoglossal axones run in linear series forward through the medulla to the anterior lateral sulcus, whence they emerge between
joins the facial nerve
the pyramid and the olive (Fig. 85). They separate the anterior from the lateral area. The main body of the anterior columna is broken up into the nucleus later alls inferior, the nucleus ambi-guus and the motor part of the nucleus of the ala cinerea.
304
THE RHOMBENCEPHALON.
The nucleus lateralis inferior (Figs. 92 and 94) is situated in the reticular substance of the lateral area. Though it is made
up
this region that the
among
naked
the fibers of
It
eye.
prob-
ably gives origin in the anterior and lateral column of the spinal cord (Tschermak). It is known to receive fibers from the anterior ascending cerebellospinal tract
cuneati,
funiculi gracilis
and
funiculi
and
the cerebellum, which probably transmits tactile, muscular, pain and temperature impressions to the cerebellar cortex.
nucleus ambiguus (Fig. 93) forms an irregular sheet of gray substance, pear-shaped in section, which extends longituIt is prolonged from dinally through two-thirds of the medulla.
The
It is
The
sory nerve run out through the posterior lateral sulcus below the level of the olive.
The nucleus of the ala cinerea (Fig. 93) is in part derived from the base of the anterior columna; this part is motor in function and its neurones resemble those of the anterior columna in
It is situated, It belongs to the ninth and tenth nerves. above, close to the floor-ependyma of the fourth ventricle under the ala cinerea; and it extends, inferiorly, into the closed medulla
the cord.
nearly as far as the hypoglossal nucleus. It is immediately lateral to the nucleus of the hypoglossal. Its axones run in a curve, convex toward the median plane, between the restiform body
olive. They are joined medially by those axones of the nucleus ambiguus which enter the roots of the ninth and tenth nerves. By this nucleus and the nucleus ambiguus, many dis-
and the
formed, belonging to the roots of the ninth, tenth and the cerebral part of the eleventh nerves. They run in slightly
tinct fascicles are
of the posterior
medulla
motor
cells of
305
cells of the terminal nucleus of the vagus nerve, which represent neurones of the posterior columna of gray matter. They receive the end-tufts of the sensory root fibers of the vagus nerve and
possibly of a small
number from
hypoglossal nucleus, the motor part of the nucleus of the ala cinerea and the nucleus ambiguus receive many fibers from the
opposite pyramidal tract, and probably from the cerebro-pontal
tracts,
The
which bring
to
inhibitory im-
longitu-
bundles
Ascending Ant.
cerebellospinal, spino-
thalamac and
rubrc spinal
tracts
Triangular
Anterior cerebello-spinal,
tract (Helwigi)
spinal tract
Fillet decussation
fillet
decussation.
(Original.)
pulses
lished
from the
cerebral cortex;
and
by
fibers of the
sensory nuclei.
The posterior columna is decapitated by the fillet. It is represented in the medulla (i) by the following terminal nuclei, viz., the sensory part of the nucleus alae cinereae of the vagus and
glossopharyngeal nerves, the vestibular and cochlear nuclei of the auditory nerve, the nucleus tractus solitarii and the nucleus
of the spinal tract of the trigeminal nerve;
and
(2)
by the gray
306
THE RHOMBENCEPHALON.
The nucleus alee cinerece of the vagus and glossopharyngeal nerves (Fig. 93) contains in its lateral part a group of small fusiform cell-bodies like those in the posterior columna. These
fusiform
cells constitute
and it is probable that a few glossopharyngeal fibers also arborize and end in the nucleus. Cortical Connection. Axones of this nucleus probably enter into the medial fillet, the spino-thalamic tract and the medial longitudinal bundle. The two former conduct tactile, muscular, pain and temperature impulses to the thalamus, whence the cortical fillet carries them to the cortex; the latter establishes its reflex connection with motor
of the vagus,
nerves.
Nucleus Tractus
Solitarii (Figs. 92
and
94).
The
nucleus of
the solitary tract surrounds the tractus solitarius with which it coincides in extent. It is gelatinous in appearance and is situated
just lateral to the nucleus of the ala cinerea.
In
its
descent
it
trends dorsally and toward the median line. It is joined to the opposite nucleus, at its spinal end, by the nucleus commissuralis The nucleus of the solitary tract is the terminal nucleus (Cajal).
of the afferent fibers of the intermediate
nerves and, probably, receives a few fibers thus the nucleus of the nerves of taste and forms the
station in the gustatory path.
relay
The axones
of the cell-bodies in
and continue the taste path toward the thalamus, but their exact course has not been worked out. Nucleus Tractus Spinalis Nervi Trigemini (Figs. 92 and 95).
ent nuclei
The nucleus
in character.
is
gelatinous
continuous with the sensory pontine nucleus of the trigeminal nerve, above; and, below, is continued in the gelatinous substance of the posterior columna of the spinal cord.
As low down as the second cervical segment it receives fibers from the trigeminal nerve, so the terminal nucleus of this nerve extends from the middle of the pons to the second cervical nerve. The nucleus of the trigeminal is embraced between the emergent part of the facial nerve, medially, and the vestibular nerve, laterally, in
307
lies
enlarges in bulk
along the ventro-medial surface of the restiform body; and approaches the surface near the middle
of
the medulla,
it is
and
rior
produces the tuberculum cinereum; then continued down into the cord as a cap of the poste-
where
it
In the lower part of the medulla of gray substance. the nucleus underlies the visible part of the tractus spinalis nervi
trigemini.
columna
The
on
its
sensory root of the trigeminal nerve (fifth) enters the pons ventral surface, in line with the roots of the seventh, eighth,
and accessory nerves (Fig. 85). The root fibers divide the short ascending branches end in the pontine nucleus T-like; of the fifth nerve and the long descending branches, forming the
ninth, tenth
certain
few
motor nucleus
of the trigemi-
nal nerve and perhaps to other motor nuclei; these are reflex in
function.
the trigeminal nucleus axones establishing reflex and cortical relations run (a) To motor nuclei by way of the medial long:
From
itudinal bundle
and
and
(b)
by way
of
The
latter
fillet
and
the spino-thalamic tract. The axones bearing impulses of the muscular sense enter the medial fillet and are continued through it
to the lateral nucleus of the thalamus; those fibers
which conduct
pain and temperature impressions run through the spino-thalamic tract to the same nucleus. Both sets of fibers conduct tactile impulses.
From
fillet
Vestibular Nuclei (Nn. Nervi Vestibularis, Figs. 92 and 93). These are located partly in the pons as already pointed out, and extend as low as the mid-medulla. Their function is equilibrium.
The
and
lies
principal micleus (Schwalbe's) is dorso-medial in position beneath the acustic area of the ventricular floor, crossed
striae.
by the medullary
It extends transversely
appears
which arbor-
308
ize
THE RHOMBENCEPHALON.
and terminate about
is
its cells.
of the descending root. the medial surface of the restispread along form body, chiefly in the pons. It becomes a distinct nucleus as the lower border of the pons is approached and grows larger
some distance above that point. In the pons it is bent backward with the restiform body toward the cerebellum. Its upper end is thus placed in the lateral wall of the fourth ventricle between the restiform body and the brachium conjunctivum. This porfor
tion
is
(of
Deiters's nucleus
made up
of cell-bodies
the
from cerebellar
ganglia and cortex and originates axones that proceed down the cord in the anterior descending cerebello-spinal tract, thus form-
ing a relay in the cerebello-spinal path; and this is its chief function. It is only a subordinate terminal nucleus of the vestibular
nerve.
nucleus of the descending root is, in all probability, a relay station in the conduction path from the vestibular nerve, but not a terminal nucleus of that nerve. It is composed of
cell-bodies scattered through a strand of fibers, called the descend-
The
ing root, which extends from the level of the principal nucleus down to the nucleus funiculi cuneati (Bruce). It is placed somewhat under cover of the medial border of the restiform body and,
with the enveloping descending root, separates this body from Certain fibers of the descending root principal nucleus. terminate in the nucleus of the same name.
the
Cortical Connections.
vestibular nerve enter the opposite medial fillet and, descending root, also the nucleus funiculi cuneati.
fillet
by way
of the
The medial
continues the direct path to the thalamus and the cortical completes it up to the temporal cortex. The descending root and the arcuate fibers from the nucleus funiculi cuneati
fillet
the
cortex
of
the
cerebellum.
The
reflex
connections of the
first, with cerebral nerves, by fibers which join the medial longitudinal bundle and terminate
309
axones of Purkinje's
bosus; from the
motor nerves; second, with spinal nerves by the cells to the nucleus fastigii and nucleus glocerebellar
by the acustico-cerebellar and then by way of the anterior tract to the motor nuclei of the spinal
ganglia
Cochlear Nuclei (Nn. Nerui Cochlearis, Fig. 92). There are two cochlear nuclei, the ventral and the lateral. They concern
hearing proper. The ventral cochlear nucleus appears in section as a triangular mass of cell-bodies imbedded hi the medulla at
the upper end of the posterior lateral sulcus. It lies between the restiform body and the olive; the vestibular root of the auditory nerve separates it from the olive. It receives the greater number
of fibers in the cochlear nerve
and
a few of
its fibers
enter the
fillet
of the
same
In the corpus trapezoideum, the cochlear tract is largely relayed by the neurones forming the nuclei of the superior olivary
group.
The
of the restiform body. It is situated both lateral and dorsal to the ventral nucleus and, stretching around the posterior surface
of the restiform body,
it
the lateral part of the acustic area, called the tuberculum acusticum. The lateral nucleus receives that part of the cochlear root which does not end in the ventral nucleus, and the fibers arborize about
its cells.
form the medullary striae; a few of them enter the trapezoid body (Figs. 86, 89 and 92). The medullary striae run somewhat obliquely across the ventricof the lateral nucleus
The axones
median groove, plunge forward to the superior nuclei of the opposite side where they are partially relayed olivary and then, bending upward, are continued in the lateral fillet.
ular floor to the
At the superior olivary nuclei of the opposite side the fibers from the lateral and ventral nuclei become intermingled, hence the
trapezoid body and medullary the lateral fillet. The lateral
striae
fillet
combine
in the formation of
own
it
310
THE RHOMBENCEPHALON.
inferius; the smaller part ends in the
brachium
quadrigeminal
side.
colliculi, chiefly in
same
From
the medial geniculate body to the transverse and superior temporal gyri, the acustic path is formed by the temporo-thalamic (acustic)
radiation.
nuclei.
established, first, by the and medial longitudinal bundle and, second, by olivary pedicle that part of the lateral fillet which ends in the colliculi of the corpora quadrigemina (see pages 160 and 161). (3) There are Certain Special Nuclei of the Medulla. These are not represented either in. the pons, above, or the spinal
Their
cord, below.
are the nucleus funiculi gracilis, the nucleus funiculi cuneati and the nucleus olivaris inferior.
They
(Figs.
The
cuneati are large nuclei, extending from the level of the olive to the lower end of the medulla. They are situated near the posterior surface
fibers terminate in
beneath the gracile and cuneate funiculi, whose them; they give origin to the medial fillet, and
the anterior and posterior external arcuate fibers, and they produce, respectively, the clava and cuneate tubercle on the posterior
In successive sections from below upward, nucleus funiculi gracilis is first seen as an isolated mass of gray substance imbedded in the funiculus gracilis at the level of the pyramidal decussation. It enlarges dorso-ventrally and toward its upper end, as is shown in consecutive sectransversely tions, and reaches its greatest size at the clava, where it receives the terminal end-tufts of the funiculus gracilis. At the level
surface of the medulla.
the
The axones
fillet
and the
(Fig. 95)
appears at the same inferior level as the nucleus funiculi gracilis. It is from the first and throughout its length continuous with the
central gray substance in the lower medulla.
on which
it
It gradually
(Fig. 94).
tubercle
reaches its full stature and gathers into itself the fibers of the funiculus cuneatus and the descending root of the vestibular nerve; thence it sends its own axones upward in the medial fillet and the
external arcuate fibers.
of the
medulla there
is
funiculi cuneati, to
a small lateral bud of gray matter connected with the nucleus which it is accessory and, like it, is imbedded
It
nucleus juniculi cuneati. The nuclei funiculi gracilis et cuneati form the
first
and
Nucleus funiculi
gracilis
Tractus spinalis
n. trigemini
Head
Ant. cerebello-spinal, spinb-thalamic, rubro-spinal triangular tracts
of ant.
columna
and
Pyramid
1
j"
Fasciculus proprius
Fig. 95.
tactile senses,
and
lesions in
them cause
fillet
ataxia.
They
also lie
the
fillet
and
the indirect path running through the arcuate fibers to the cerebellar cortex. From the cerebellar cortex the impulses proceed
312
THE RHOMBENCEPHALON.
and, thence, through the dentate neurones by way of the brachium conjunctivum cercbelli to the opposite red nucleus and thalamus.
The
cortical
fillet
conducts
all
common
The
94), is a sinuous, pouch-like collection of gray matter rethe nucleus dentatus of the cerebellum. It is situated sembling near the lateral surface of the medulla and is invested superficially
(Figs. 92
and
and deeply by
which join
it
fibers
Its
open
with
to the opposite
On
the
an accessory nucleus
medial accessory, in the anterior area among the fibers of the interolivary part of the medial fillet, and the dorsal accessory in.
the lateral area.
lateral fasciculus
The
by
fibers of the
(oliva).
The
olive
shows the longitudinal extent of the nucleus and on section it is seen to measure a quarter of an inch in depth. The olivary nucleus is said to be a modern structure; it is found well developed only
in the higher mammals and does not appear in the human brain It probably contains both ascending and until the sixth month. neurones. Axones enter it from the nucleus funiculi descending
gracilis
and nucleus
funiculi cuneati
of the
receives
cere-
two groups
The descending
down from
the hilus of the opposite nucleus, arborize and end about its cellbodies; and (b) the olivary fasciculus (central tract of the tegmentum), having descended from the lentiform nucleus through the mid-brain and pons, fades
away
and
probably ends hi it. Axones of the olivary nuclei of the medulla have been traced by Kolliker and others down the lateral column
of the spinal cord.
The
composed
of such fibers.
cord lateral to the anterior roots of the spinal nerves and ends in the gray matter of the cord.
313
Lesions in the medulla are very fatal and death usually occurs before any sensory or motor phenomena can be observed; but rarely the pyramidal tracts alone have been involved or the pyramidal tracts together with
In the last case, crossed paralysis as in the pons, affecting the cerebral nerves on the produced, same side and the opposite spinal nerves. In progressive bulbar paralysis the motor nuclei of the medulla are involved as a preliminary to the degeneration of the anterior gray columna in the
spinal cord.
RHOMBENCEPHALON.
SECTION
The common
is
IV.
cavity of the
the* fourth
118).
The
fourth ventricle
contained chiefly in the pons and medulla, and is ventral to the cerebellum (Fig. 79). It is broadest at the junction of the pons and medulla (Figs. 86 and 91). Above and below that junction,
it
central canal of the spinal cord, with which it is continuous. Inferiorly it communicates through its roof with the subarachnoid
space via three apertures, a median and two lateral. The fourth ventricle is a gable-roofed chamber with a diamond- shaped
floor.
The
gables
(Fig.
79)
and are
prolonged in tunnel-like extensions around the restiform body. The long axis of the ventricular floor (Figs. 86 and 96) is parallel with the spinal cord, and extends from the superior extremity of
the pons to the middle of the medulla. The transverse axis Thus the coincides with the junction of the pons and medulla.
superior triangle of the floor by the medulla oblongata.
is
inferior,
The
lined with
ependyma, which
is
the inferior part, where below the inferior medullary velum only the epithelial layer is present.
Boundaries.
The
floor
is
314
THE RHOMBENCEPHALON.
The
lateral wall (superior triangle) is formed by the brachium conjunctivum of the cerebellum; and (inferior triangle) by the
taenia of the fourth ventricle
funiculus cuneatus and funiculus gracilis to the obex. The roof is formed by the superior medullary velum (valve of Vieussens) superiorly; and by the inferior medullary velum and roof epithelium, inferiorly (Fig. 91). The superior and inferior halves of the roof meet at an acute angle, the fastigium, and form the tent On either side, the gable is of the fourth ventricle (Fig. 79).
is
formed the
lateral
The
lateral recess is
The ular cavity, reaching almost to the posterior lateral sulcus. and ventrally, by the restiform body;
by the
inferior
inferiorly,
by the
roof epithelium. The chorioid plexuses of the fourth ventricle invaginate the roof epithelium and hang from the roof into the
lateral recesses
and the inferior part of the cavity (Fig. 91). Floor of the Fourth Ventricle. (Fossa rhomboidea). Because it contains the nuclei of one or more roots of the posterior
eight (fifth to twelfth) cerebral nerves, the floor of the fourth ven-
A median groove tricle is a very important area (Figs. 86 and 96). bounded by the eminentiae mediales forms the long axis of the diamond-shaped floor and divides it into two lateral halves; the lateral halves are bisected transversely by a number of lines, the medullary striae (striae medullares). The striae are produced by bundles of fibers which rise from the cochlear nucleus of the Diverging somewhat and plunging into the auditory nerve. medulla at the median groove, the fibers of the striae enter the opposite trapezoid body and lateral fillet. The medullary striae divide each lateral half of the floor into a superior and an inferior
triangle.
Description to Fig. 96.
Nucleus of olfactory nerves, b. Nucleus of oculomotor nerve, c. Nucleus of trochlear nerve, d. Nucleus of descending root of trigeminus. e. Chief motor nucleus of trigeminus. f. Nucleus of facial, g. Nucleus of abducens. h. Nucleus ambiguus (vagus and glossopharyngeus). i. Nucleus of (hypoglossus. j. Nucleus of accessory nerve. Nuclei of optic nerve: k. Pulvinar of thalamus, 1. Lateral geniculate body, m. Nucleus of superior colliculus. n. Sensory nucleus of trigeminus. o. Nucleus of vestibular nerve, p. Ventral nucleus of cochlear nerve, q. Lateral nucleus of cochlear nerve, r. Nucleus alaecinereae (vagus and s. Solitary tract (vagus and glossopharyngeus) t. Nucleus of spinal glossopharyngeus)
a.
.
.
tract of trigeminus.
315
-P
96.
and
Nuclei of the cerebral nerves in the medulla, pons, mid-brain, inter-brain Motor (or genetic) nuclei, red; terminal (or sensory) olfactory bulb. nuclei, blue. (After Morris's Anatomy.)
317
The
coiliculus
facialis
is
(Fig.
86),
superior
extremity
it is produced largely by the genu of the nucleus of the abducent (sixth) nerve (Fig. 96). External to it and in front of the striae medullares is a small fossa, the fovea
It is
superior.
Fovea Superior
(seventh)
(Fig. 86).
The
fovea superior
is
and the
salivary nucleus
which are deeply seated in the pons. Running upward along the wall of the ventricle from the superior fovea, is the sulcus limitans. It is a blue-floored groove in the pons, called locus
caeruleus.
The
of the ventricle.
The
blue color
is
due
nea, a pigmented layer of cell-bodies underlying the ependyma. The principle motor nucleus of the trigeminal nerve (fifth) lies
beneath the superior part of the locus caeruleus (Fig. 96). Inferior Triangle of the Ventricular Floor. It presents:
The trigonum
cinerea,
and eminentia
and most
The hypoglossal
of the ventricle,
its
apex
base looks upward and is situated under the medullary striae. The twelfth nerve rises from the column of cells whose upper onehalf
is
covered by
it
(Fig. 96).
glossi
and
which forms the apex of the ala cinerea. Ala Cinerea (Trigonum Vagi, Fig. 86).
is
The vagus
it,
triangle
hence the
name, ala
cinerea. The inferior fovea forms the depressed and directed apex of the ala; its floor rises inferiorly to the superiorly The base, eminenlia cinerea, which is directed toward the clava.
318
nucleus
alas
THE RHOMBENCEPHALON.
cinereae, the
nucleus tractus
solitarii
cleus ambiguus, three nearly parallel columns of cell-bodies a halfinch in length, are in part covered by the ala cinerea (Fig. 96).
The
cus,
fossae
sul-
of the
Area Postrema.
Below the
and between
it
and the
a small fusiform strip of the venAn tricular floor which Retzius has called the area postrema.
taenia ventriculi quarti, there is
oblique stria separates it from the base of the ala cinerea. The area acustica occupies the lateral angle of the ventricular
floor (Fig. 86).
It is partly in the superior triangle,
but chiefly
in the inferior.
its
of the acustic area most promBeneath the acustic area are the vestibular nuclei of the auditory nerve; also the lateral part of the cochlear nucleus, which
is
(Fig. 96).
intermedius
The
first,
six of
and
twelfth,
are purely motor; while the. fifth, the intermediate, the ninth and tenth are mixed nerves and contain both efferent and afferent
fibers.
Cerebral Nerves, Nervi Cerebrales (Figs. 86 and 96). 1. Olfactory (nn. olfactorii) special sense of smell.
2.
Optic
(n.
opticus)
(n.
special
sense
of
sight.
3.
Oculomotor
Trochlear
4.
(n.
319
Trigeminal
(n.
6.
7.
Abducent
Facial
(n.
(n. facialis)
Intermediate
secretory and
8.
special
sense
of
taste,
trophic.
special senses of hearing
Acustic
(n. acusticus)
and equisense of
librium.
9.
Glossopharyngeal
taste,
(n.
glossopharyngeus)
Special
common
(n.
and motor.
10.
Vagus
vagus)
n. Accessory
12.
Hypoglossal
hypoglossus)
AU
when
their
mixed nerves such use of the term "origin" is not admissible. Pure sensory nerves and the sensory roots of mixed nerves take
from ganglia situated wholly outside the brain. From those ganglia the dendrites grow outward to the peripheral distribution of the respective nerves; the axones grow centrally into
their origins
the brain, where they arborize and end in groups of cell-bodies forming nuclei. Such nerves conduct impulses from the per-
iphery to these nuclei, hence the name applied to them is terminal nuclei (nuclei terminates). See the blue nuclei, Fig. 96. The motor nerves and the motor roots of mixed nerves take their
origins inside the brain
nuclei.
from groups
the periphery; they conduct impulses from the nuclei to the muscles or to the secreting cells hi their respective areas of distribution,
hence the nuclei of motor nerves and motor roots are genetic nuclei (nuclei origines). See the red nuclei, Fig. 96. Thus it is seen
that the brain connection of a
motor nerve
is its
a sensory nerve.
320
THE RHOMBEXCEPHALON.
TABLE
H.
Terminal Nucleus.
Olfactory (Smell).
Olfactory
cells
in
nasal
Under
surface of olfac-
Mitral
and brush
cells
mucous membrane.
tory bulb.
bral hemisphere).
2.
Optic (Sight).
Nuclei of same in interbrain and mid-brain.
perior quadrigeminal
colliculus.
5.
Semilunar ganglion.
trigemini
reaching
ond
cervical nerve.
Geniculate ganglion.
Nucleus tractus
in
solitarii
and
eighth
8.
Cochlear Root.
Spiral
Cochlear Nuclei.
tral
Venplaced
in
and
lateral,
ventral
and
lateral to
restiform
dulla.
body
me-
Vestibular Root.
Vestib-
ular ganglion.
and nu-
cleus of descending
root, in floor of fourth
ventricle in medulla.
9.
etc.}.
Superior
ganglia
and
in
Nucleus tractus
solitarii
alae
jugular
and nucleus
ereae in
cin-
foramen.
medulla.
321
Vagus (Sensory
Root}.
Jugular and
nodular
in jugular
it.
Nucleus
alae cinereae,
and
ganglia
medulla.
TABLE m.
Nucleus
ulus.
in floor of cerebral
in mid-brain
4.
Nucleus in
ulus.
floor of cerebral
aqueduct
in mid-brain
under
inferior collie-
rhombencephali.
5.
Nucleus
in
uleus of pons.
6.
in
7.
Nucleus
alae cinereae
upper end.
Vagus (Motor
Root).
Same
ambiguus.
and between
body.
olive
and restiform
322
THE RHOMBENCEPHALON.
ii.
Cerebral Root.
Nucleus ambiguus
closed medulla.
below
level of olive.
poste-
segments of cord.
12.
fourth ventricle,
and
in
olive,
closed medulla.
Terminal Nuclei.
The
first
and second
cerebral nerves are peculiar and cannot as yet be classified with the nuclei of other sensory nerves and sensory roots (Figs. 26, 43 and 63). The terminal nuclei of the fifth, the intermediate,
the eighth, ninth and tenth nerves may be called the posterior columna series; because they are formed by masses of cell-bodies
representing the
upward prolongation
of the posterior
columna
mon sensory and special sensory. Common Sensory Nuclei (Fig. 96).
the posterior columna series of nuclei, the terminal nucleus of the fifth and of the tenth
Of
nerves and a part of the terminal nucleus of the ninth nerve receive common sensory impulses, and transmit them to the opposite thala-
mus
by two routes,
viz.,
spino-thalamic the former route; those of the pain and temperature senses, over the latter; and tactile impressions are believed to traverse both
routes alike.
tract.
through the medial fillet and through the Impulses of the muscular sense travel over
From the thalamus these impulses' are carried to the cortex of the posterior central gyrus. Thus is the cortical connection of these nuclei established; and each is brought into
reflex connection
nuclei which run chiefly through the medial longitudinal bundle and terminate hi the motor nuclei.
The
impulses from the glossopharyngeal and intermediate nerves, has not been definitely traced but it is probably relayed in the thalamus and is established by
which receives
323
certain fibers of the internal capsule which end hi the fusiform gyrus (?). The cochlear nuclei (ventral and
in the hemisphere,
by
impulses of hearing and conduct them on toward the cerebral cortex by way of their axones, which form the trapezoid body and medullary striae and then unite in forming
fillet
the lateral
(Fig. 89).
The remaining
connection are formed by the brachium inferius and the acustic The reflex connection radiation (radiatio temporothalamica).
of these nuclei
is
somewhat
indirect.
fillet
It is established in part
by
which end
in the quadrigeminal
chiefly
brought
together with the anterior longitudinal bundle; but is about by the olivary pedicle and the me-
dial
The -vestibular nuclei the principal longitudinal bundle. Deiters's and the nucleus of the descending root (Schwalbe's),
They
receive impulses
concern equilibrium.
pleted
by
the cortical
by the opposite medial fillet, and comfillet; and the latter is formed by the de-
scending root, which ends in the nucleus funiculi cuneati, and the With the cerebral motor nerves, these external arcuate fibers.
first, by axones entering the medial longitudinal bundle and terminating in the motor nuclei; and, second, probably by fibers of the brachium conjunc-
third, fourth
and
The
reflex
nerves
viz.,
by axones of Purkinje's
the cerebello-olivary fibers and descending fibers from Deiters's and the inferior oh" vary nuclei.
The
geniculate body, the pulvinar of the thalamus and the superior colliculus of the corpora quadrigemina (Fig. 43). Like the ter-
minal nucleus of the olfactory nerve, these cannot at present be included in the posterior columna series, because the ventral and
dorsal zones of the embryonic fore-brain have not been sufficiently
324
elucidated.
THE RHOMBENCEPHALON.
If the sulcus
hypothalamicus really separate ventral from dorsal zone in the inter-brain, as is claimed by many, it would
optic
included in the posterior series but there is need of further investigation, as this places the whole cerebral hemisphere in the dorsal zone.
The
by
body and
in the pulvinar of the thalamus and terminate hi the cortex of the calcarine region of the occipital lobe. From this cortical center,
corticifugal
fibers
few fibers directly from the outer root of the optic tract; it thus becomes the center of optic reflexes; and axones of the superior colliculus form the anterior longitudinal
colliculus also receives a
bundle, which completes the connection with opposite motor nuclei. The terminal nucleus of the olfactory nerve is situated in the olfactory bulb (Figs. 26
and
63).
The axones
by way
first,
with the uncinate region and anof the lateral olfactory stria
;
and, probably, by way of the intermediate stria and, second, with the parolf actory area, the olfactory triangle, the gyrus subcallosus and the gyrus cinguli through the fibers of the medial stria of the
olfactory tract.
The
nerve
is
:
but
following probabilities First, the fornix, stria medullaris thalami, fasciculus retroflexus
of interpeduncular ganglion; second, the fornix, thalamo-mammillary bundle and descending axones of the thalamus;
little
understood.
The
may be
stated
and axones
and, third, the fornix and the pedunculo-mammillary bundles running into mid-brain, pons and medulla.
(Fig. 96).
The
nuclei of the
oculomotor, trochlear, abducent, facial, sal nerves and the nuclei of the motor roots of the trigeminal, glossopharyngeal and vagus nerves represent the anterior columna
of gray matter in the cord
series.
and constitute the anterior columna These nuclei are connected "with the cerebral cortex on both
325
The
con-
nection
established
some
and
fillet
of the fibers
first and principally by the pyramidal tracts, running directly from the tract to the nucleus,
(Bechterewi) to a point
run through the accessory near the respective nuclei which they
are
about
to
enter;
and, second, the fronto-pontal, temporotracts are believed to send some fibers to
the genetic nuclei of the same side. The reflex connection of these genetic nuclei is established for all of them by the medial longitudinal bundle; by the anterior longitudinal bundle and by the
olivary pedicle
(for the
third, fourth,
and
sixth),
by the
trapetri-
zoid
body (for the seventh) and by the spinal tract of the geminal nerve (for the fifth, seventh, and twelfth).
CHAPTER
V.
Dura Mater.
Through
the foramen
magnum
the
membranes
which they are very similar in structure. The dura mater spinalis is 'attached to the margin of the great foramen and to the bodies of the first
of the brain are continuous with those of the cord with
cervical vertebrae; elsewhere, though joined to the by fibrous bands, its surface is free from immediate bony attachment and it does not possess the periosteal layer. Thus
vertebrae
two or three
suspended, it hangs as an open sack, or sheath (Fig. 100) and reaches down to the third sacral vertebra, where it is constricted
to a fibrous cord
which blends with the periosteum on the posterior The arachnoid and pia, and the spinal
cord and cauda equina are contained in the dural sack (Figs. 97, 98 and 100). Externally, the surface of the dura is separated from the wall of the spinal canal by the internal vertebral plexus
of veins, areolar tissue
and
fat.
The
outer surface
is
composed
surface
bathed with a small amount of cerebro-spinal fluid which For every segment of the spinal separates it from the arachnoid. cord, the dura presents, on either side, a pair of foramina, through
is
Those nerve roots are invested by a sheath of dura (Fig. 97). from the margins of the foramina. The dura mater prolonged of the cord does not separate into two layers, and forms neither sinuses nor processes. It performs no periosteal function and no arachnoid granulations (pacchionian bodies). Its possesses two surfaces are formed by endothelium. Arachnoid. The arachnoid of the spinal cord (arachnoidea spinalis) forms a sack of the same length as the dural sheath, with which it is externally in contact (Figs. 97, 98 and 100). It
326
327
Septum posticum
Dura mater
Ligamentum
denticulatum
Anterior nerve-roots
(in sections)
Subarachnoid space
Dura mater
Arachnoid
Posterior root
Anterior root
Ligamentum denticulatum
Linea splendent
Fig.
97.
Meninges
A. Transverse section. of the spinal cord. (After Morris's Anatomy.) (After Ellis.
Key and
3 29
presents two serous surfaces. Internally, bands of nbro-elastic tissue attach it to the pia mater along the posterior median line of the cord and form the subarachnoid septum (Fig. 97). The
external spinal veins
and a considerable space separate the arachThat subarachnoid space is filled with
it is
By
and
same spaces in the cranial and 6). cavity (Figs. 97 Pia Mater. The pia of the cord (pia mater spinalis] is much stronger than that of the brain (Figs. 97 and 98). It has two
magnum,
Ligamentum denticulatum
Body
of
vertebra
Periosteum
Dura mater
Subdural space
Arachnoid Subarachnoid
space Pia mater
Intervertebral foramen
Fig. 98.-
-Diagrammatic section of the spinal meninges and spinal cord. (After Morris's Anatomy.}
which is continuous with the brain and forms an epineurium for the cord and roots of the spinal pia The outer is the more vascular layer. Both layers dip nerves.
distinct layers, the inner of
they form the anterior septum which contains the anterior spinal artery. The inner layer is
fissure;
median
attached to the septum in the posterior median fissure. The outer layer forms the linea splendens along the front of the cord, and the
330
ligamenlum denticulatum on either side. The denticulate ligament is a longitudinal band whose smooth medial border is continuous with the pia along the middle of the lateral surface of the cord; its lateral border is notched and its twenty teeth, invested
vertebrae.
with arachnoid, are attached to the dura opposite the first twenty The two ligaments subdivide the space between the
pia and arachnoid into anterior and posterior subarachnoid spaces. filamentous extension of the pia below the cord proper helps
form the filum terminate internum. It descends in the arachnodural sheath with the roots of the lumbar and sacral nerves, and
to
together constitute the cauda equina (Fig. 100). For some distance, about half its length, the filum terminale internum conall
and rudimentary
fibers
spinal cord.
The
the third sacral vertebra in forming the filum terminale externum which forms a sort of ligament fbr^he spinal cord. That ligament is inserted into the coccyx. The "pia mater of the cord contains the trunks
and large branches of the anterior and the two and the tributaries of the external spinal
Nerve Supply. The membranes of the spinal cord are supplied by recurrent branches of the spinal nerves and by the sympathetic.
The
and the two posterior spinal arteries, which rise at the foramen magnum from the vertebral arteries, and are reinforced by cervical,
intercostal
arteries. The anterior spinal artery descends along the entrance to the anterior (a. spinalis anterior) median fissure (Fig. 99) it is formed by the union of two vessels,
and lumbar
(a.
spinalis
freely
which
The
is
331
The spinal arteries give origin to two sets of branches, the fissural or centrifugal, and, the centripetal arteries. namely, Both sets are end-arteries and form rich longitudinal plexuses,
which overlap each other but do not anastomose. The fissural or centrifugal arteries rise, first and chiefly, from the anterior spinal artery (Fig. 100). These enter the
anterior
median
fissure
and,
Fig. 99.
The
arteries
Diagrammatic.
a. Dorsal external spinal veins, b. Posterior radicular vein. c. Peripheral venous plexus. d. Anterior radicular vein. e. Ventral external spinal veins, f. Anterior central vein. g. Posterior central artery and vein. h. Posterior spinal artery, i. Peripheral arterial plexus. k. Intercostal artery. 1. Spinal ramus. m. Anterior radicj. Posterior radicular artery, ular artery, n. Interior spinal vein. o. Anterior central artery, p. Anterior spinal artery.
greater part of the gray matter. Second, a few centrifugal arteries rise from the posterior spinal arteries. Running into the posterior
fissure,
posterior
they are distributed to the posterior white columns, the commissure and to the nucleus dorsalis (Clarki).
The centripetal
angles to the surface,
arteries
rise
They
332
Those branches
Veins.
to the columnar
The
veins that
the cord, the venae spinales internae, are the fissural veins, which issue from the fissures, the root-veins, which accompany the anterior
and posterior
and a
small
number
(venae spinalis externae) spread over the entire surface of the cord beneath the arachnoid membrane. According to Cunning-
anterior
either side,
an
antero-lateral
and and a
In the upper cervical region, the plexus forms two or three small veins which empty into the vertebral or inferior cerebellar veins;
by a branch along each spinal nerve, the plexus communicates with the internal vertebral plexus (plexus venosi vertebrales intern!) outside the dura mater, and is drained into
elsewhere,
No
valves
perineural
spaces
carry
the
vessels
CHAPTER
VI.
and forms the corresponding portion of the central axis of the nervous system. Extent. It is continuous with the medulla oblongata, above; and, in the adult, reaches to the lower border of the first lumbar
part of the neural tube,
vertebra (Fig. 100). Its length is seventeen to eighteen inches. In a very slender process, the filum terminate internum, the cord
is
continued beyond the first lumbar vertebra. That process and the lower spinal nerves form the cauda equina, which is inclosed in a sheath composed of the arachnoid and dura mater.
The
prolongation of the central gray matter and ventricle of the cord; and, also, a few fibers, which suggest the coccygeal nerves of lower
animals.
In the foetus before the third month, the cord and spinal canal are of equal length. At birth the cord reaches the third lumbar vertebra, and it continues to recede with the rapid growth of
the vertebras to adult
life.
Diameters
(Fig. 101).
The
spinal cord
is
shaped
like
a cylinIts
(dorso-ventrally).
less
longest diameter
is
transverse
and measures
except hi the cervical and lumbar enlargements of the cord. In the latter, it equals a half -inch; and, hi the former, it slightly
exceeds
it.
The
cylindrical in shape.
thoracic portion of the cord is small and nearly Divested of its meninges and nerves the
and a
half, avoirdupois.
The
cervical
enlargement
(intumescentia
cervicalis)
extends
from the medulla oblongata to the second thoracic vertebra Its greatest diameter is on a level with the (Figs. 100 and 101).
fifth intervertebral disk.
It gives origin to
333
334
and brachial
plexuses.
enlargement (intumescentia lumbalis] begins at the tenth thoracic vertebra and increases to the twelfth (Figs.
The lumbar
Opposite the
first
lumbar vertebra,
it
tapers off
imtim*mr-^*
of Cord.
/uiv
Inferior Portion of
Cord and
Cauda Equina.
Posterior view of the spinal cord, the dura mater and the arachnoid being laid open and turned aside. (Brubaker after Sappey.) Floor of fourth ventricle. 2. Brachium conjunctivum. 3. Brachium pontis. 4. Res9, 9, 9, 9. n, n, II, n. 13. Anterior roots of spinal 15. Conus medullaris. 16, 16. 8.
5. Clava. 6. Glossopharyngeal nerve. 7. Vagus. 10, 10, 10, 10. Posterior roots of Posterior lateral sulcus. 12, 12, 12, 12. Spinal ganglia. 13, nerves. 14. Anterior and posterior divisions of spinal nerve. Filum terminales internum. 17, 17. Cauda equina. I-VHI. racic nerves. I-V. Lumbar nerves. I-V. Sacral nerves.
tiform body.
Accessory nerve.
Ligamentum denticulatum.
spinal nerves,
Cervical nerves.
I-Xn. Tho-
almost to a point, the conus medullaris; but a very small process continues in the filum terminate internum. From the lumbar
enlargement rise the motor fibers lumbar and sacral plexuses, and of the same plexuses.
into
it
335
median
sept.
Postero-lat. sul
Gelatinous substance
Fig. 101.
The
cervical, B.
The
thoracic, C.
The
(Original.) A. Section of cervical cord. A. C. Anterior columna, P. C. Posterior columna, C. G. Gray commissure, anterior gray and posterior. Com. A. White anterior commissure. Fun. Ant. Funiculus anterior, Fun.' Lat. Funiculus lateralis. Fun. Post. Funiculus posterior. G. Fasciculus gracilis, C. Fasciculus cuneatus. B. Section of thoracic cord. L. C. Lateral columna. C. Section lumbar cord. D. Section of lower sacral cord.
lumbar, and D.
The
sacral.
337
The
cord (canalis centralis spinalis) is the representative of the cavity It is just visible to the naked eye; but it of the neural tube.
ventricle.
extends throughout the cord and expands above into the fourth In the filum terminale internum it is also dilated,
It is lined with forming the ventriculus terminalis (Krausei). columnar ciliated cells which stand on a thick lamina of substantia
gelatinosa.
is
The spinal cord (Fig. 101). divided into symmetrical lateral halves by the incompletely anterior and the posterior median fissure.
fissure
(fissura
mediana
It
anterior) is
extends in
length from the inferior end of the ventral surface of the pons (foramen caecum of Vicq d'Azyr) down the anterior median line
of the
As
to depth,
it
formed by the white anterior commissure. Both layers of pia mater dip down into it and inclose the anterior The anterior median fissure is spinal artery and its branches.
cord's axis.
interrupted at the junction of the cord and medulla by the decussation of the pyramids. In the lumbar enlargement it gradually
disappears.
is
fissure (fissura mediana posterior) narrow and deep (Fig. 101). It extends, longitudinally, down the posterior median line of the cord from the middle of the
posterior surface of the medulla.
ventrally,
It
dorso-
beyond
its
middle.
The
by the
median
The
posterior
median
fissure is not
an open
fissure; occupied by a lamina of connective tissue, the posterior septum, which is attached to the deep layer of the pia mater. In the posterior septum ramify branches of the two posterior spinal
arteries
and
PosteriorLateral Sulcus.
half of the spinal cord
is
Each lateral
partially divided, near the junction of the posterior fourth with the anterior three-fourths of its semi-
It
and the
antero-lateral surface
from each
it is
Anterior Root-line.
lateralis anterior).
As a landmark,
the most lateral fibers of the anterior roots of the spinal nerves, the anterior root-line of the spinal cord (Fig. 101). There is no
groove on the surface of the cord along this line and it is misleading to call it a sulcus, as has been the custom. It is situated opposite
the anterior columna of gray matter and in line with the anteriorlateral groove of the medulla oblongata. Through it and through the surface, just medial to it, emerge the anterior roots of the
spinal nerves.
rior
It
and
lateral surfaces.
(s.
intermedius posterior)
a slight longitudinal groove in the cord which subdivides the upper three-fourths of the posterior surface into postero-medial surface and postero-lateral surface (Fig. 101). From it a connective tissue
ciculus gracilis
septum extends into the cord and separates the fasand fasciculus cuneatus from each other. The
is
The
spinal
cord
composed
grisea spinalis), in the central part; and (2) white matter (substantia alba spinalis) in the peripheral area. It is like the medulla
and pons in having the white matter on the surface A column of gray matter (Fig. 101), crescentic
(Fig. 101).
in section,
extends through the center of each lateral half of the spinal cord. The crescent is convex medially; and is joined to its fellow, a
little
by a
vertical transverse
lamina of
339
anterior commissure.
white matter of the opposite side by the white The points of the crescent are directed
lateral projection
columna in the cervical and Together, the two crescents and the
of gray matter.
The
H-shaped column is well marked in the cervical and thoracic regions; but, toward the lower end of the cord, the crescents become short and thick, and the gray column is almost cylindrical.
The H-shaped column is composed of two kinds of gray substance, viz.: (i) The substantia gelatinosa (Rolandi), which forms
a cap for the head of the posterior columna and (b) an en(2) The velope for the central canal, or ventricle, of the cord.
(a)
substantia spongiosa.
The
latter
forms
all
the H-shaped
column
except the tips of the posterior columnae and the thick sheath of the central canal. Imbedded in the neuroglia, there is a network
of medullated nerve fibers;
and
common
stains,
give rise to
Gray Crescent
columna;
(2) the
made up
to
which
lateral
joined
fellow of the
lateral
opposite side
projection,
called
columna; and,
is
(3) the
posterior
not everywhere clear cut and definite, especially in the cervical region, but is intermingled for a short distance with the white matter, forming the
lateral
columna.
The
formatio reticularis.
The
formatio reticularis
it
is
found in the
(Fig. 101) as
is
seen in sections
short
columna.
It is thickest
and thick compared with the posterior in the cervical and lumbar enlargements,
toward the
it
where
it
is
more
slender.
the surface of the cord as does the posterior columna. It ends in a bulbous, serrated head, which points toward the anterior
root-line,
and
is
by the
cervix
340
or base.
From it the anterior roots of the spinal nerves rise; and, with the anterior root-fibers, it separates from each other together the anterior and lateral white columns of the cord.
The gray Cells of the Anterior Columna (Figs. 102 and 103). matter of the spinal cord contains multipolar neurones of the
Golgi and Deiters types. The Golgi cells ramify richly in the gray matter about the cell-bodies, and, both their axones and
dendrites terminate in relation with other neurones in the adjacent gray substance. The long axones of the Deiters cells either enter
into the anterior roots (radices anterior),
called radicular
cells, or they enter into a longitudinal tract or strand of fibers, and the neurones are named strand-cells (Cunningham). The dendrites of the Deiters cells arborize in both the
The cell-bodies in the anterior columna and vesicular in character. They are motor or efferent function, and their axones form, in great part, the anterior roots
of the spinal nerves. Together with the neurones of the genetic nuclei of cerebral nerves, these of the anterior columna constitute
Two
chief
columns
of cell-bodies
column and
column
(Figs.
102 and 103). The former is continuous throughout the cord with the exception of the fifth lumbar and the first sacral segments
(Bruce- Cunningham); while the lateral column
in the cervical of cells
is
found only
shows a double group in sections of the lower three cerand the first lumbar segments of the cord.
called
dorso-
Only the ventro-medial group is present above the sixth cervical segment and below the first lumbar segment. The
dendrites of the cell-bodies in the medial
column arborize
in the
gray substance of the same columna, in the adjacent white matter of the anterior column of the cord and, to some extent, in the
opposite anterior columna, having passed through the white anterior commissure; the axones of these medial cell-bodies enter
very largely into the anterior roots of the spinal nerves on the same side; but a certain number probably run through the white
341
Mid- cervical
Fasciculus gracilis Fasciculus cuneatus
Gelatinous substance
Entry zone
Marginal tract
(Post.) Cerebellospinal fasciculus
Dorso-lateral
cells
Triangular tract
Antero-lat. fascic ulus proprius
Ventro-medial
cells.
Mid-thoracic
Comma tract
Lateral 'pyramidal tract
Gelatinous substance
Rubro-spinal tract-
-4
Nucleus dorsalis
(Clarki)
Descending ant.
cerebello-spinal fasciculus
Intermedio-lateral
cells
Dorso-medial
Antero-lat. fasciculus proprius
cells
Ventro-medial
cells
Tracts of 'fibers -and columns^of cells, in the cervical and thoracic regions Fig. 102. o'f'the cord. Diagrammatic. (In part after Bruce and Cunningham.)
343
commissure into the anterior nerve-roots of the opposide, and others enter into the fasciculi proprii of the cord. The lateral column of cells in the anterior columna is a large
(Figs. 102
one
and
It is
103).
It is
found only in the regions which is, in the cervical and lumbar
enlargements.
dorso-lateral cell-group,
and
in
enlargement there are two other cell-groups, according to Alexander Bruce. One of them is located behind the dorso-lateral cells
and
is
and the
other,
which
occupies the angle between the ventro-lateral and the dorso-lateral cells, lying medial to both, is called the central group (Cunning-
ham).
arborize
The
columna
and
proceed largely into the anterior roots of the spinal nerves but partly into the longitudinal white columns of the cord. Probably the medial column innervates the trunk muscles; the lateral column,
the
muscles
of
the
extremities.
Cortical Connection.
These
cell-columns are brought into relation with the anterior (direct) pyramidal fibers and the lateral (crossed) pyramidal fibers by
means
of
intermediate neurones.
In
this
inhibitory impulses descend to them from the cerebral cortex, coming from the opposite hemisphere, chiefly, but also from the same side. They also are in relation with the end-tufts of posterior
and with axones whose cell-bodies are located in the center and posterior columna of the gray crescent. The latter neurones form contact relations with fibers of the posterior roots
root-fibers
on both
sides.
contact between anterior and posterior root-neurones and by the intervention of an intrinsic spinal neurone the simple reflex mechanism of the spinal cord is formed. It has been the belief that the
end-tufts of the fibers in the anterior
and
lateral
pyramidal tracts
are in. direct contact with the dendrites or cell-bodies of the neu-
rones in the anterior columna; but the investigations of Schafer, Collier and others, indicate that this connection between the
fibers,
344
at least,
is established by intervening neurones whose cell-bodies are located near the base of the posterior columna in the region of the nucleus dorsalis (Clark's column). The evidence of such
termination of the anterior pyramidal tract Lesions. Paralysis due to lesions of the
is
not conclusive.
anterior
columna
seat
and
of
is
segment paralysis.
The
hemorrhagic inflammation and rapidly degenerate in acute anterior poliomyelitis. In progressive muscular atrophy and
in amyotrophic lateral sclerosis they degenerate slowly.
result of the
first,
As a
sudden
The muscles
waste away in the second and third because the nerves controlling the muscles and their blood supply are gradually destroyed. In the last, the muscles are also spastic, because the involvement
of the pyramidal tracts cuts off cerebral inhibition.
(2)
(Fig. 101).
such
as,
the cilio-spinal, cardia'c-accelerator, vaso-motor, secrecell-bodies in the center of the crescent are of
;
That the
sympathetic function is suggested by two facts first, the cell-bodies are small, which indicates that the axones run but a short dis-
thetic
tance from the neurone center, as is the case with spinal sympaneurones; and, second, the distribution of these central
is
neurones
whence
B)
is
The
intermedia-lateral
column
of cell-bodies
(Fig.
102,
the only one found in the center of the crescent. In the thoracic segments of the cord, where the lateral columna is visible, this
column
is
immediately adjacent to
so far as
it
is
and lumbar enlargements it is situated in the base of the anterior columna near its lateral surface. The intermedio-lateral column is found hi the last cervical, all the thoracic and the first and second lumbar segments, in a nearly continuous column; it is also found
345
column
Entry zone
Marginal tract
(Lissaueri)
Central cells
Triangular tract
(Helwigi)
Ventro-lateral cells
Ventro-medial
cells
Dorso - latera
cells
Central cells
Descending ant.
cerebello-spinal tract
Tracts of fibers and columns of cells in the lumbar and sacral regions Fig. 103. of the cord. Diagrammatic. (In part after Bruce and Cunningham.}
347
and fourth
and
in the
The
first
lumbar segment) corresponds in position and extent to the origins of the white rami communicantes the second region is at the level of origin of the pelvic splanchnics; and the
cervical to second
;
cell-groups representing this column in the upper cervical segments probably contribute sympathetic fibers to the accessory
The
cells of
the intermedio-lateral
column are
largely of the
radicular variety, their slender axones enter into the anterior Whether any belong to the strandroots of the spinal nerves.
variety
is
not known.
They
root-fibers
and are
by
intervening neurones.
traced.
(3)
lower cord,
columna and where it receives the posterior roots of the spinal nerves. The posterior columna presents a slight enlargement near its extremthe caput columnce, which tapers off to the apex ity, called columns. The head is joined to the base by a constricted part,
the cervix.
posterior] except in the slender (Fig. 101). It is longer than the anterior reaches the surface in the posterior lateral sulcus,
The head
of the posterior
columna
is
capped by a
V-shaped mass of substantia gelatinosa. Spongy substance makes up the remainder of it. The posterior columna separates the
posterior
The
of of
column of the cord (Figs. 102 and 103). columna are numerous. In the head the posterior columna they have smaller bodies than the cells the columna anterior. They are less definitely grouped and
from the
lateral
cells of the posterior
are fusiform in shape throughout the caput columnar, but in the base of the horn, near its medial surface, they have large vesicular bodies and form one of the most definite cell-columns in the
spinal cord.
posterior sensory in function.
The
afferent or
The
number
largely to the type of Golgi, as the processes of of them neither enter into the anterior roots
348
of the spinal nerves, nor help to form the longitudinal fiber-tracts of the cord. Their axones arborize and end in the gray substance
Those axones of the Golgi cells which cross over through the posterior commissure to the opposite crescent help to constitute the "immediate decussation" of the pain and
of both crescents.
temperature path. There are also Deiters cells in the head of the posterior columna; their axones enter into the posterior fasciculus proprius of the cord, hence they belong to the strand-variety.
They appear
of the
to
field of
and
at least a part
comma,
oval and septo-marginal tract. The neurones columna receive posterior root-fibers
which carry excito-reflex impulses and impulses of the tactile, pain and temperature senses. The former are transmitted for-
anterior
columna
the latter, to a considerable extent, are carried through the posterior commissure to the anterior columna of the opposite crescent.
Nucleus Dorsalis (Stillingi and Clarki). This column, which was discovered by Stilling, is composed of cell-bodies measuring from 40 to 90 in diameter (Figs. 102 and 104). It forms a
fj.
It is situated
bounded laterally by a curved strand of posand forms a continuous column from the seventh cervical segment to the second lumbar segment. The column is largest in the lower two thoracic segments, where it bulges out
posterior columna,
terior root-fibers ;
the medial surface of the posterior columna. It is represented by separated groups of cell-bodies in the third and fourth sacral
and
three or four cervical segments of the cord and in the accessory nucleus of the cuneate column in the medulla oblongata. The limitation of the dorsal nucleus, as an unbroken column,
first
rami communicantes has suggested its connection with the sympathetic system; and it is the terminal nu-
cleus of afferent sympathetic fibers, but it gives rise to no efferent fibers of that system. All the axones of the dorsal nucleus appear to
and enter
349
same
side.
The
Together
with the cell-bodies, the dendrites are in contact relation with fibers of the posterior roots of the spinal nerves. The nucleus
dorsalis probably has to
do with muscular
tonicity
and equilibrium,
Concerning the relation of the gray crescent to the spinal nerves (Fig. 104), it may be remarked, here, that in the anterior columna
Posterior
median
fissure
Posterior root
Fibers to medulla
Anterior root
Fig.
104.
The
Diagrammatic.
(Original.)
Anterior root rises from medial, lateral and intermedio-lateral cells of same crescent and from medial cells of opposite crescent: it is also connected with the opposite side by certain dendrites. Posterior root terminates in gelatinous substance, in the center and base of the posterior columna, in the intermedio-lateral column, and in the anterior columna of the same crescent in the anterior and posterior columna of the opposite side ( ?) and in the nucleus f uniculi gra cilis or nucleus funiculi cuneati of the medulla oblongata.
; ;
and center of the crescent are located the genetic nuclei of the motor or efferent fibers (anterior roots) of the spinal nerves;
columna, but also in the center and anterior columna of the crescent in the
cord,
funiculi gracilis
and nucleus
funiculi
350
The gray commissure of the spinal cord (the gray anterior, and the posterior commissure) is the vertical, transverse sheet of gray substance connecting the two crescents together (Fig. 101). This commissure (commissura grisea) completes the gray matter
It unites the gray crescents together a little in front of their center, except in the lumbar region where it joins their It forms the floor of the posterior median fissure; and, centers.
of the cord.
in front,
is
is
It
which
pierced longitudinally by the central canal of the spinal cord, is surrounded by a thick envelope of substantia gelatinosa.
ventricle, divides the
That part
of the
the
gray anter'ior commissure (commissura anterior grisea) and that behind it is the posterior commissure (commissura posterior, Fig. The gray commissure, comprising both these divisions, 101).
is
are
of
composed of spongy and gelatinous substance in which there imbedded the bodies of many nerve cells and a large number medullated fibers. The medullated fibers are derived from
the posterior roots of the spinal nerves, and from intrinsic neurones of the cord, whose centers are situated chiefly in the commis-
sure and in the posterior columna. The posterior commissure is said to contain a long sensory tract, between the ventricle and
dorsal surface (Ciaglinski). This long sensory tract is found in the thoracic portion of the cord and the discoverer believes it to
be made up of ascending
temperature impulses.
(Barker).
It
root-fibers
is
in
Lesions of the gray substance, as in syringomyelia, completely abolish the pain and temperature senses at the level of the lesion, while the muscular and tactile senses are preserved. The dissociation of sensations is most complete and bilateral when the
commissure; if the lesion be limited one crescent, the pain and temperature senses are affected on the side opposite to the lesion. These facts show that the pain
lesion destroys the posterior
to
to the cerebral cortex run through the gray substance of the cord and that they decussate, through it imme-
cord.
It also
351
impressions may reach the cerebrum without passing through the gray substance in the spinal cord; but it does not exclude the possibility of a part of the impulses of the muscular and tactile
senses being transferred from a lower to a higher neurone in the spinal gray substance.
2.
white matter (Fig. 101) of the spinal cord (substantia alba spinalis} is disposed in its peripheral area and in the white anteIt is composed of medullated nerve fibers and collaterals) imbedded in a small amount of neuroglia, (axones and supported by a connective tissue network derived from the
The
rior
commissure.
pia mater.
vessels.
it is
The
and
ventrally
The transverse
in
tracts into the gray
tracts; (2) the
which are usually somewhat oblique those running from the longitudinal
matter or out of the gray matter into such axones of intrinsic neurones which connect the two
same level; and (3) posterior root-fibers the posterior commissure to the opposite crescent. running through The white anterior commissure of the spinal cord (commissura
anterior alba} is the only definite lamina of transverse fibers in the cord (Fig. 101). It connects the anterior and lateral white columns of the cord with the opposite gray crescent and the two
gray an-
composed
commissure, forming the floor of the anterior median fissure. of medullated fibers belonging to (a) the anterior
tract,
pyramidal
and spino^thalamic tracts; (d) it the crossed fibers to the anterior roots of the spinal comprises nerves, and (e) the decussating dendrites between the anterior
cending anterior cerebello-spinal
columnae.
fibers of the spinal cord (Fig. 104) are those of the anterior roots of the spinal nerves, in their course (a) from the gray matter to the surface of the cord; (b) those of the
The dorso-ventral
352
posterior roots, running from the posterior-lateral sulcus to their destination in the gray matter, and (c) axones of intrinsic neu-
rones connecting posterior with anterior parts of the crescent. The longitudinal fibers comprise most of the white matter
in the cord,
anterior,
funiculus lateralis
and funiculus posterior (Fig. 101). These three great columns occupy the anterior, lateral and posterior areas of the cord. They are disposed around the gray crescent in bundles or tracts. The tracts which make up the funiculi are not visible to the naked eye, nor under the microscope in a healthy adult cord; they have been located by embryological, experimental and pathological
investigations.
The
spinal cord
and
longitudinal fibers rise in the brain, hi the some run upward and others
tracts of the cord.
Thus
the tracts
are characterized as ascending, descending and mixed tracts. Ascending Tracts: (i) Ascending anterior cerebello-spinal
and spino-thalamic
bellar)
tract.
(2)
with the spino-vestibular tract. (3) Marginal tract (of Lissauer). (6) (4) Fasciculus cuneatus. (5) Fasciculus gracilis.
Ciaglinski's long sensory tract, hi the posterior commissure.
Descending Tracts:
pyramidal
tract.
(i)
including Helwig's
tract.
Rubro-spinal
called
tract.
tract
(Pawlow).
levels
Descending postero-medial
tract,
at
(6)
different
the
comma,
oval,
and septo-marginal
Descending postero(i)
lateral tract.
The
ante-
proprius,
Lateral fasciculus proprius. (3) The posterior fasciculus composed of the cornu-commissural tract, the septal
bandalette of Hoche, and possibly a part of the comma, oval, septo-marginal and the descending postero-lateral tract (Cun-
ningham).
The methods
may be summarized
briefly, as follows:
first
employed successfully
353
by Flechsig. He found that nerve fibers when first laid down are naked axones without any insulating white substance of Schwann ensheathing them. That the medullary sheaths are developed at different times and that the medullation is nearly coincident
with the beginning of function. Thus the fibers of motor and sensory nerves are first to become medullated, since life cannot be sustained without the automatic mechanism. Second, the fasciculi proprii of
automatic and coordinating mechFourth, the voluntary motor mechanism is established by the medullation of the tracts connecting the lower neurones with the cerebral cortex, the fibers of the pyramtracts.
At
idal
tracts
last
This
being the last to receive their medullary sheaths. occurs just before birth. Fibers of the cerebrum con-
become medis
ullated gradually, year after year, keeping pace with the mental
not com-
(Kaes).
The pathological and experimental methods depend upon the fact that a nerve fiber when severed from the cell-body undergoes degeneration in accordance with the law of Waller. If the severed fiber be above the cell-body, the degeneration occurs
above the lesion and is called ascending degeneration; but, if the degeneration extends from the lesion down the nerve fiber, the cell-body being above, then the condition is called descending
degeneration,
though
all
parts
of
the
severed
fibers
really
the paths of dedegenerate simultaneously. generation, above and below a destructive lesion hi the human cerebro-spinal axis, the various tracts of fibers have been discov-
Thus by studying
ered and
ation.
many
of
origin to termin-
These
investigations
study of degenerations in the brain and cord of lower animals. These degenerations are the results of definite experimental lesions,
as cutting of certain posterior nerve roots, partial section, hemisection or complete section of the spinal cord, etc. The pathoand experimental methods are commonly called the physiological
logical method.
23
354
(fasciculus antero-
occupies the deep part of the anterior and lateral columns (Figs. 102 and 103). It embraces the anterior columna of gray matter and covers the outer surface of the center of the
crescent
By
the most
of the cord.
fissure
approaches, but does not quite reach, the surface Notice that it is separated from the anterior median
tract,
Behind,
it is
composed and descending fibers which are the T-branches of axones from the gray crescent. It is largely a short fiber tract, associative and commissural in function. That part situated in
of ascending
The
antero-lateral
fasciculus proprius
the anterior column, the anterior fasciculus proprius, is largely commissural, between the anterior columna:; while the lateral
fasciculus proprius
is
chiefly associative,
and connects
different
segments of the cord on the same side. The antero-lateral fasciculus proprius is continued in the substantia reticularis of the
medulla,
and the
reticular
constituting a short fiber tract which extends from the lower part of the cord to the basal ganglia of the cerebrum. In the anterior
fasciculus proprius there
dial longitudinal
is a strand of long fibers called the mebundle (fasciculus longitudinalis medialis).
.
(posterior] longitudinal bundle is composed of an and a descending strand of fibers (Figs. 102 and 103). ascending The ascending strand rises from the anterior columna in each
The medial
segment of the spinal cord and runs upward to the motor nuclei of cerebral nerves and terminates in them. Perhaps a few fibers
reach the thalamus.
It
columna in the cord; in the medulla, it runs between the head of the anterior columna and the pyramidal decussation, then just
lateral to the
fillet
it
takes
its
dorso-medial
The descending
355
It rises from is the medial panto-spinal tract (Collieri). the nuclei of the reticular formation but chiefly from the nu-
and the nucleus lateralis medius in the pons. Forma part of the medial longitudinal bundle of the same side, ing its fibers end in the center of the crescent as it descends the cord.
clei centrales
Within the
descends the
lateral ponto-
decussates.
It
origin right through the medial longitudinal bundles. Both ponto-spinal tracts extend to the lower part of the cord.
tract
(fasciculus cerebro-spinalis
anterior), occupies a thin area next the anterior median fissure It is the direct continuation of about 10 per (Figs. 1 02 and 103). Its fibers are axones of the pyramidal tract in the medulla. cent,
of cortical cells
gyrus of the cerebrum. As the tract descends in the cord, the fibers decussate through the white anterior commissure, and ter-
cells of
the
posterior columna.
the opposite gray crescent, probIt reaches to the fifth sacral the anterior pyramidal tract
segment
is
(Collier).
Imbedded
in
a small strand
first
bundle.
Anterior Longitudinal Bundle (Figs. 102 and 103). Held called It occupies a very narit the fasciculus longitudinalis ventralis. row strip in the anterior column just beside the entrance of the
anterior
median
fissure.
The
its
through the dorsal tegmented decussation (Meynerti) to a position in the mid-brain ventro-lateral from the medial
It descends in that relative position through longitudinal bundle. the pons and half the medulla; near the pyramidal decussation
the anterior and medial longitudinal bundles are brought together and lie between that decussation and the isolated head of the an-
columna; they diverge upon entering the cord and remain separate to the end. The anterior longitudinal bundle ends in both anterior columnae (Collier). It forms the middle link in
terior
the ocular
and pupillary
reflex arcs.
356
Loewenthal's column) together with the ascending anterior cerebello-spinal tract, occupies a thin peripheral area, broadest poste-
which extends from the anterior pyramidal tract outward and backward, over the antero-lateral fasciculus proprius, to the middle of the lateral surface of the cord (Figs. 102 and 103). Its
riorly,
posterior border
tract
is
and the
lateral
pyramidal
tract.
The
fibers of the
two an-
scending fibers are found, chiefly, in the anterior and posteromedial part of the common area and the ascending in the posterolateral part.
dulla.
almost completely relayed in the mefibers, in the first stage, are axones from
is
fastigii
the cortical cells of Purkinje in the cerebellum and from the nucleus and perhaps the other cerebellar ganglia. They descend
to the nucleus of Deiters
cerebelli.
There the greater number end and new fibers rise which continue the tract down the cord. In all probability the cerebello-olivary
fibers in the restiform
fibers,
forming
the tract of Helwig in the cord, should be included in the descending cerebellar tract. It terminates in the gray substance of the
spinal cord
and
neurones
is
by the intervention of intrinsic spinal connected with the motor cells of the anterior coldirectly or
umna. of an
The descending
and
anterior
columna neurones,
it
reflex
and equilibrium.
Scattered
the vestibolu- spinal fibers in the posterior part of the anterior descending cerebello-spinal tract are descending fibers from the thalamus and quadrigeminal colliculi which end in the
among
gray crescent.
Ascending Anterior Cerebello-spinal and Spino-thalamic Tract. (Fasciculus ascendens cerebello-spinalis anterior, Gowersi). These form a single tract in the cord which is found chiefly in the
postero-lateral part of the area
common
to
it
and
103).
composed
357
and base
opposite side of the cord (v. Lenhossek). Most of the fibers of the ascending anterior cerebello-spinal and spino-thalamic tract
cross near their origin through the white anterior
commissure of
the cord.
tract ascends
part of the lateral area, sending collaterals to the inferior lateral nucleus; it then continues, through the formatio reticularis of the
pons, to a point near the inferior quadrigeminal colliculus, where the cerebellar part is bent backward under the brachium conjunc-
tivum, and enters the vermis cerebelli superior through the superior medullary velum (Hoche). The spino-thalamic portion continues to the thalamus, sending
colliculi
some
fibers to the
quadrigeminal
and lentiform
nucleus
The
anterior
ascending
pain,
cerebello-spinal
and spino-thalamic
impulses.
is
tract
carries
tract
tactile,
and temperature
The
triangular
of
Helwig
(fasciculus olivaris)
common
It
to the
descends
to the
lumbar
It probably rises region, where its longest fibers end. in the olive of the medulla oblongata and is efferent in conduction.
The
(posterior)
cerebello-spinal
[posterior])
tract
(direct
cerebellar)
(fasciculus
cerebello-spinalis
runs
posterior
to
the
other cerebello-spinal tracts (Fig. 102). It is superficially located, and in section extends from the middle of the lateral surface of
the cord back to the posterior lateral sulcus as far down as the lumbar cord. Below the second lumbar segment its absence allows the lateral pyramidal tract to
(posterior)
come
to the surface.
The
cerebello-spinal tract
worm
of the cerebellum.
axones of vesicular
cells hi that
nucleus.
In the
forms a part of the restiform body. It conveys impulses medulla, of the muscular sense, received, especially, from the viscera (?).
tract is
a small
strand of fibers discovered by Horsley and Thiele in 1901, called the spino-vestibular tract. It rises in the lumbo-sacral region
358
to the restiform
body
The
lateralis}
lateral
pyramidal
tract
(fasciculus
cerebro-spinalis
forms a considerable part of the lateral column of the cord (Figs. 102 and 103). It is covered, superficially, by spinal the cerebeUo-spinal tracts in the cervical and thoracic cord; but
in the
lumbar and
sacral cord
it
Its
is in relation with the posterior columna of gray matter, the lateral fasciculus proprius and the marginal bundle. The fibers composing it are axones of cell-bodies in the anterior
deep surface
They
rise
anterior pyramidal tract, and the two run as one tract down through the genu and anterior two-thirds of the occipital part of the internal capsule, the middle three-fifths of the basis pedun-
the anterior longitudinal fibers of the pons and the pyramid of the medulla. In the medulla the two tracts separate. The
culi,
lateral
tract,
with
its
median
fissure,
pierces the
columna and descends with some uncrossed fibers column of the cord. It terminates in relation with
the cell-bodies within the posterior columna, according to Schafer, Collier and others. The anterior tract follows the anterior median
fissure as already described.
Both end
According to Marchi, ten or twenty per cent, of the fibers remain uncrossed. The pyramidal tracts are the cerebral motor tracts. By them motor and inhibitory
impulses are carried to the cord. In the outer part of the lateral pyramidal area is found the crossed descending tract of the red nucleus, the rubro-spinal tract (Pawlow).
the center of the gray crescent. Its origin in the nucleus ruber, its crossing through the ventral tegmental decussation (Foreli) and its course down the brain
102).
It
The
rubro-spinal tract
is
descending
359
the
lateral)
The pyramidal
tracts
(especially
are
involved in lateral sclerosis and in amyotrophic lateral sclerosis; and, as a consequence of it, both voluntary and inhibitor}' impulses
from the brain are interfered with, hence the spastic paralysis and exaggerated reflexes. The pyramidal tract may be more or less involved in insular sclerosis and in bulbar paralysis, and
the
the
pyramidal, cerebello-spinal
which
affects
is
legs.
In ataxic paraplegia
and posterior the degeneration in the pyramidal tracts that causes the spastic gait, incoordinated arm movements and early increase of the reflexes, observed in that affection.
(Gowersi) there
diffuse sclerosis of the lateral
It is
The marginal tract (fasciculus marginalis, Lissaueri) is a small tract composed of the ascending branches from the outer set of fibers in the posterior roots of the spinal nerves (Figs. 102
and
103).
It is situated in
and apex
and, after ascending a short distance, end about the cell-bodies of the substantia gelatinosa.
Column
of the
Cord
(Fig. 101).
In the posterior column of the spinal cord, there are, first, two ascending and two descending tracts, derived from posterior nerve roots; second, the posterior fasciculus proprius, which is
much
scattered
and
is little
occupied by the incoming fibers of the posterior roots. Entry Zone (Figs. 102 and 103). Over the apex
and along
the medial surface of the posterior columna of gray substance the posterior roots of the spinal nerves enter the cord and divide
T-like
into
is
ascending
The name
well applied to this region. zontal fibers distinguishes the entry zone
tracts.
entry zone
The
presence of hori-
The
root-fibers
of
very soon enter the gray substance. The large fibers, in part, enter the dorsal nucleus, but the greater number form the longit-
360
udinal
tracts
begin their
root-fibers
The ascending -fibers posterior column. course in the entry zone. As they ascend they upward are crowded toward the median septum by the entrance of new
the
succeeding upper segments. So the fibers near the septum are those which enter low down in the cord and those close to the posterior columna are of recent entrance into
in
the
the
cord.
The
descending
T-branches and
collaterals
of
the
Some
course, even
cervical to the
greater number are much shorter. are crowded in a dorso-medial directhey tion, like the ascending fibers; and, after a considerable downward course, they plunge forward through the white column to end in the posterior columna. These ascending and descending fibers,
The
whose origin
is
on the
proprius which rise in the posterior columna, constitute the longitudinal tracts of the posterior column of the spinal cord. That
posterior
column
is
thoracic segment, where the posterior intermediate furrow and septum fade away; but, above that level, it is subdivided into
two
distinct
gracilis,
ascending tracts, a postero-medial, the fasciculus and a postero-lateral, the fasciculus cuneatus (Fig. 102).
tracts are alike in constitution.
These two
They have
the
same
muscular and
tactile senses.
They differ only in length; the fibers of the fasciculus gracilis come from the spinal nerves below the eighth thoracic, while those of the fasciculus cuneatus come entirely from thoracic and cervical
nerves.
Fasciculus Gracilis (Ascending Postero-medial Tract, Goll's Column). This tract may be said to begin at the entrance of the It posterior root of the coccygeal nerve (Figs. 102 and 103).
ascends along the posterior median septum to the nucleus funiculi gracilis of the medulla. Up to the lower thoracic nerves it gradually enlarges,
fibers;
due
only the entry zone and the descending branches of the posterior roots, it comprises the whole
and in
361
column.
Its size is
uniform in the
upper three-fourths of the cord, where the posterior intermediate furrow separates it from the fasciculus cuneatus. In depth it
branches of the spinal ganglia
funiculi gracilis.
ascending Arriving at the clava, all the fibers of the fasciculus gracilis arborize and end in the nucleus
cells.
Fasciculus
Cuneatus
(Ascending
Postero-lateral
Tract,
Burdach's Column).
Beginning in the lower thoracic segments (Fig. 102) the fasciculus cuneatus ascends between the entry zone
fasciculus gracilis to the nucleus funiculi cuneati of the
It acquires
and the
medulla.
its
from every spinal nerve above In origin and grows stronger up to the first cervical nerve.
fibers
new
section
of the
wedge-shaped, being broadest at the surface the edge wedge touches the junction of the posterior columna and
it is
;
Its fibers are ascending, and are branches posterior commissure. of the axones of spinal ganglia cells, like the fasciculus gracilis.
arborize
and terminate
The
from the posterior roots of the spinal nerves descending themselves somewhat roughly into two tracts, a posteroarrange These descendlateral and a postero-medial (Figs. 102 and 103).
ing radicular tracts greatly expand and multiply the terminal relations of the posterior root-fibers. Mingled with them, there
are
proprius.
(Figs. 102
and 103)
tract (of
levels.
In the cervical
Its ventral part
is
the
comma
disappears in the posterior columna above the tenth segment, but the remainder continues down the cord. Shifting its position
in a dorso-medial direction,
it
takes
its
septum and gradually approaches the cord's posterior surface. It continues in that situation to the end of the cord, and is called,
in succession, the
oval tract
septo-marginal
tract
median triangular
362
(of
postero-medial descending tract in the cervical, thoracic, and, at least, the upper lumbar segments. After a variable course within the tract, the fibers plunge forward into the posterior
this
columna of gray substance where they terminate. The descending postero-lateral tract (Fig.
at the posterior surface of the cord
103), situated
medial to the entry zone, in the lumbo-sacral region, appears to have been first described by Thiele and Horsley. It is very largely endogenous, but contains
a number of root-fibers (Collier).
Its fibers terminate in the posterior
In section
it
is
triangular.
columna
of gray substance.
Both the descending radicular tracts are intermingled with endogenous fibers that belong to the fasciculus proprius. Posterior Fasciculus Proprius. (Fasciculus posterior proprius}.
This
is
made up of
It is
scattered throughout the posterior column, but they are especially numerous in the descending radicular tracts, in the region along
the ventral one-third of the septum (the bandelette of Hoche) and in the cornu commissural field of Marie (Figs. 102 and 103).
The cornu commissural tract (Fig. 103) is placed between the posterior columna (cornu), the posterior commissure and the It extends up to the eleventh thoracic segment posterior septum.
and downward to the end of the cord. It contains both ascending and descending fibers as do other parts of the fasciculus proprius.
muscular and
If the entry
Lesions in the posterior columns cause disturbances of the tactile senses, and ataxia and incoordination result.
zone
is
involved there
is
disturbance of
all
kinds of
common
sensation,
These columns
are usually involved, by extension from the posterior roots, in locomotor ataxia (posterior sclerosis), hence the paraesthesia,
crises, loss of reflexes,
363
These roots descend more or less from their (Figs. 100 and 104). cord attachment to the inter- vertebral foramen in which they
unite to form the spinal nerve.
The
roots of the
first
cervical
nerve are horizontal; those of the first thoracic nerve descend the width of two vertebrae, and those of the twelfth thoracic, the width
of four vertebrae; while the roots of the coccygeal nerve extend from the first lumbar vertebra to the second piece of the coccyx,
all
spinal nerves,
is
composed of from four to six fasciculi, which soon combine into two bundles. After piercing the dura mater, the anterior root unites with the posterior, beyond the latter' s ganglion, and forms
a spinal nerve.
tion.
The
anterior root
is
efferent,
or motor, in func-
Apparent Origin
(Fig.
of medullated axones
The anterior root is composed 104). which issue from the narrow longitudinal
area at the junction of the anterior one-fourth with the posterior three-fourths of the cord's surface. This area is bounded laterally
by the anterior
root-line,
commonly
sulcus.
rise
from
on the same side of the cord and from the medial column of the
opposite side. These cell-bodies of the anterior columnae and the intermedio-lateral column constitute the genetic nuclei (nuclei
origines) of the spinal nerves.
The
from the
They are voluntary motor fibers. In the intermedio-lateral column, the small fibers of the anterior roots take their origin. They are
probably sympathetic in junction, that is, involuntary motor, vaso -motor, viscero-motor, inhibitory, secretory, trophic, inhibitosecretory,
and
inhibito-trophic.
(spinal and cerebral) are probably in a state of toxic irritation in laryngismus stridulus, tetanus, acute ascending paralysis (Landry), strychnine poisoning,
Lesions.
364
etc.,
the twitchings, spasms and convulsions. Their deIn struction causes flaccid paralysis (lower segment paralysis).
spinal meningitis both the anterior
and posterior
is
(radix posterior]
It is
composed of from six to eight fasciculi, which also combine at once into two bundles. The posterior root pierces the dura mater separately from the anterior root. It unites with the anterior root in the Near the outer end, it presents a swellintervertebral foramen. which contains large vesicular unipolar cell-bodies and is called ing
first
cervical nerve;
and
is
s pinole).
The
The
posterior root are occasionally absent posterior root, external to the ganglion,
made up
These
distant parts of the body; they are the sensory fibers of the spinal
nerves.
is
On
composed
of axones,
the axonic and dendritic processes are medullated. Apparent Central Termination (Fig. 104). The posterior roots of the spinal nerves enter the posterior-lateral sulcus; and, at
once, divide into an outer set of small fibers
large fibers
and an inner
set of
with some small ones interspersed. The fibers of each set bifurcate into a large ascending and a smaller descending
branch.
branches.
Collaterals rise
Real Central Termination, Terminal Nuclei (Fig. 104). The ascending divisions of the outer set of fibers run a short distance
along the external surface of the posterior columna, and end in ramifications about the cell-bodies of the substantia gelatinosa.
They form
the marginal tract (of Lissauer). Probably their debranches have the same ending. The T-branches and scending collaterals of the inner set of fibers from the posterior root run
(i)
To
all
parts of the
and
to the anterior
side,
365
and columnae
These
those several regions of the crescent at various levels: (a) At the same level as the nerve, (b) at a lower level than the nerve, through
the descending radicular tracts, and (c) at a higher level, through the collaterals given off by the ascending tracts. (2) The large ascending T-branches of the inner set of fibers run to the me-
gracilis
and fasciculus
They terminate in the nuclei funiculi gracilis and cuHence the terminal nuclei (nn. terminales) of any spinal
level,
same
and
divide the fibers of the posterior roots of the spinal nerves into four groups, viz.: i. Excito-reflex 2. Fibers fibers which terminate in all parts of both crescents.
Physiologically, we
may
tactile senses
from muscles,
tendons, joints skin, through the ^posterior column, to the nucleus funiculi gracilis and nucleus funiculi cuneati. 3. Fibers
carrying impulses of the muscular sense from the viscera to the dorsal nucleus. 4. Fibers transmitting pain, temperature and tactile impulses to that part of the same and opposite crescent
and
cerebello-spinal
The posterior roots of the spinal nerves and the spinal are affected in locomotor ataxia, and the lesion extends ganglia to the marginal tract (of Lissauer) and the posterior column of the cord. Excepting the fasciculus proprius, the whole posterior
Lesions.
CHAPTER
VII.
TRACING OF IMPULSES.
Having studied the grouping and chaining together of neurones, let us now make the knowledge practical by tracing impulses through the better known paths formed by these various neurone groups. The paths thus formed are of three kinds, namely: I. Efferent, or motor. II. Afferent, or sensory general and special sense.
III. Reflex.
I.
The CEREBRO-SPINAL (Fig. 105) are direct, as they do not pass through the cerebellum. Their impulses ultimately run either through the spinal or the cerebral nerves,
and are both motor and inhibitory. Hence the increased reflexes and spastic contractions of lateral sclerosis in which these tracts
are diseased.
Starting in the (Fig. 105). three-fourths of the gyrus centralis anterior of the cerebral upper cortex, motor and inhibitory impulses run down through the corona
i.
OR PYRAMIDAL PATHS
radiata, the anterior two-thirds of the occipital part of the internal capsule, the middle three-fifths of the basis pedunculi, the anterior longitudinal fibers of the pons, and the pyramid of the
lateral
and anside
pyramidal
tracts to the
same
but chiefly in the opposite side of the spinal cord. By the former route, the impulses cross over in the medulla, through the decussation of the pyramids, and descend in the lateral column of the spinal cord to the gray substance in the vicinity of the nucleus
where the path is relayed, and intrinsic neurones carry the impulses forward into the anterior columna; but by the anterior route, they descend in the anterior column of the cord and decussate, in succession, through the white anterior comdorsalis (Clarki),
366
367
Accessary
fillet
(Bechterewi)
Nucleus pontis
Medulla
Pyramid
Anterior
Pyramidal decussation
Lateral pyramidal tract
pyramidal tract
Cord
105.
Direct motor paths from cerebral cortex, to cerebral and spinal nerves.
Diagrammatic.
(Original.}
Motor paths extending from the cortex of the anterior central gyrus to the nuclei of the motor cerebral nerves and of the anterior roots of the spinal nerves represents point where the section is pierced by a longitudinal fiber; 3, 4, 5, 6, 7, 9, 10, n, and 12, nuclei of
;
.
cerebral nerves.
369
Impulses by either route finally reach the anterior gray columna of the spinal cord and, with the exception of a small per cent, of them, they reach the columna opposite to their cortical The few undecussated fibers in the lateral pyramidal origin.
conduct uncrossed impulses to the anterior columna of the same side. Thus are explained two symptoms of hemiplegia due to cerebral lesion, viz., weakness on the well side and slight motion on the paralyzed side. From the anterior gray columna of the spinal cord the nerve commotions are conducted by the efferent, or motor fibers of the spinal nerves to the muscles. 2. Through the Cerebral Nerves (Fig. 105). Impulses destined to the cerebral nerves run chiefly from the lower two-fourths
tract
genu of inand on, by the same path as the impulses to spinal nerves, down to the point where they leave the pyramidal tract to enter the nuclei of the cerebral nerves, which some of them do
ternal capsule
in the vicinity of the several nuclei.
These fibers leave the pyramidal near the internal capsule, and descend through the medial portion of the fillet to points near the respective nuclei in which
they end by multiple division. From either hemisphere impulses proceed to the nuclei of both sides. But the greater number
enter the nucleus of the fourth nerve on the
nuclei of the third,
fifth,
same
side,
and the
By
the above
orbit; the
and
and
styloid muscles;
the muscles of the larynx, trachea and bronchi, and of the pharynx, esophagus, stomach, and the intestines down to the descending
colon.
And
vasodilator,
trophic
THE CEREBRO-PONTAL PATHS, FRONTAL, TEMPORAL AND INTERMEDIATE. These paths are indirect, for
spinal nerves, since they run through the cerebellum.
24
370
TRACING OF IMPULSES.
(Fig.
Fronto-pontal
frontal
106).
The impulses
originate
in the
region
(?),
and descend through corona radiata, the and the medial fifth of the
Temporo-pontal.
They
the temporal cortex and run through corona radiata, the occipital part and inferior lamina of the internal capsule, and the
lateral fifth of the basis peduncli to the nucleus pontis.
Both
the fronto-pontal and the temporo-pontal paths are probably relayed in the corpus striatum or the thalamus. Intermediate
Bundle.
Cortical impulses of
unknown
by
the corpus striatum and conveyed, by a bundle of centrifugal axones which form the deep portion of the basis pedunculi, to the substantia nigra and then to the nucleus pontis, chiefly of the
same
side.
The impulses
broad but thin area hi the basis pedunculi just ventral to the substantia nigra and dorsal to the pyramidal tract. In all three of
these paths the impulses run to the nucleus pontis of the same side and to motor nuclei of the cerebral nerves. Whence they
(i)
To Spinal Nerves.
They run
down the anterior descending cerebello-spinal tract to the anterior gray columna of the spinal cord. Their course from the cerebellar cortex is through the acustico-cerebellar tract and
the restiform body, the lateral area of the medulla and the anterolateral column of the cord. From the gray matter of the spinal
cord the impulses are conveyed by the motor fibers of the spinal nerves to the muscles which they supply. (2) To Cerebral Nerves.
The
by the
the cerebro-pontal tracts, through the motor fibers of these nerves to their distribution.
of
(Fig. 106).run from some Impulses part of the cerebral cortex to the thalamus and red nucleus or to
and an
indirect.
the corpus striatum and red nucleus. From the red nucleus they pursue either a direct or indirect route.
(i)
route,
Direct Route, The Rubro-spinal Path. By the direct impulses run through the crossed descending tract of the
371
Temporo-pontal tract
Fibers from red nucleus to^nucleus dentatus
and
spinal tract
Rubro-spinal tract
Cord
Anterior root of spinal nerve
Fig. 106.
Diagrammatic. (Original.)
spinal nerves. A. Frontopontal tract rising in frontal lobe. B. Intermediate tract rising in lentif orm nucleus. C. Temporo-pontal tract rising in middle and inferior temporal gyri. Also rubro-spinal tract. represent points of perforation in the sections.
373
red nucleus to the center of the gray crescent in the opposite side of the spinal cord. Crossing the median raphe at once, in the
hypothalamic region through the ventral tegmental decussation, (Foreli), the impulses descend by way of the rubro-spinal tract,
through the ventral part of the formatio reticularis of the midbrain and pons, in the medial part of the lateral fillet, then through
the lateral area of the medulla,
among
and
finally down the spinal cord, the ventral portion of the lateral pyramidal area, to their through destination in the gray matter. From the center of the gray cres-
ascending
cerebello-spinal tract,
cent they proceed with or without transferring to the roots of the spinal nerves, and are conducted to the muscles.
v
(2)
Indirect Route.
may
also
opposite
down
the
anterior
which continues
columna anterior
ponto-spinal
of the cord.
tract
is
PONTO-SPINAL PATHS.
name
suggested by
The
the
The medial ponto-spinal tract accompanies the medial longitudinal bundle down the anterior ground bundle of the cord without decussating, unless the crossing occurs near the
either side.
termination.
The
lateral ponto-spinal tract, which is crossed, its origins and descends in the
Just
column
how
and nuclei
w hich
r
the ponto-spinal tracts take their origins, cannot be having arrived in them, they descend to
all
both crescents of the spinal cord and apparently enter into its segments. The anterior nerve roots complete the paths.
chiefly,
(i)
Impulses having reached the great ganglia of may run through many relays down
the formatio reticularis of mid-brain, pons and medulla and the antero-lateral fasciculus proprius of the spinal cord, to the gray
374
crescent of the same,
TRACING OF IMPULSES.
and continue through the anterior
root-fibers
to their destination. On the other hand, the impulses, leaving formatio reticularis in mid-brain, pons or medulla, may enter the nuclei of motor cerebral nerves and be conducted by them to
the muscles and glands supplied by cerebral nerves. (2) The the pons, and run impulses may leave the formatio reticularis in to the cerebellar cortex through the brachium pontis. From the
cerebellum they
may
follow the ordinary course through the anby way of the nucleus
columna
of the cord.
(3)
Impul-
by way
of the brachia
conjunctiva and brachia pontis, may descend through the cerebello-olivary tract, in the restiform body, to the opposite inferior
olive,
of the olive,
column, to the gray matter of the cord (Kolliker). (4) Impulses descend from the nucleus of the third cerebral nerve by way of certain fibers in the medial longitudinal bundle to the
in the lateral
seventh nerve, where it enters into the colliculus facialis, and through the facial nerve innervate the following muscles the
frontalis, procerus, corrugator
and orbicularis
oculi.
Hence
these
muscles are not paralyzed in nuclear facial paralysis. But if the lesion be in the colliculus fascialis or distal to it, then facial paralysis is complete. (5) At the same eminence, impulses pass into the facial nerve from the hypoglossal nucleus via a small strand of axones which rises in the hypoglossal nucleus and runs through
the medial longitudinal bundle into the facial at the genu (?). These impulses innervate the orbicularis oris. (6) Through
in the nucleus of the sixth cerebral nerve
certain short fibers in the medial longitudinal bundle which rise and cross to the opposite
nucleus of the motor oculi, impulses run from the nucleus of the abducent through the opposite third nerve to the internal rectus
oculi.
They
also the conjugate deviation observed in lesions affecting the nucleus of the sixth nerve. In nuclear lesions of the abducent
and
nerve the external rectus of the same eye and the internal rectus if the nucleus is destroyed and stimulated if the nucleus is only irritated.
375
Red nucleus
Dscussation of brachia conjunctiva
Brachiuni. conjunctivum
Vestibular root
VHIjnerve
Medial
fillet
IX nerve
Nucleus funiculi gracilis Nucleus funiculi cuneati
Medulla
Thoracic nerve
Fig. 107.
Common
column and
in the
377
sensory paths conduct two varieties of impulses, viz., general and special. The impulses originate in the end-organs of the cerebral and spinal nerves, and by those nerves are con-
The
veyed to the cerebro-spinal axis, through which they reach the proper cortical area in the cerebrum.
i.
General Sensations.
General sensation
is
This
the tactile sense, mussense has four important subdivisions cular sense, pain sense, and temperature sense. Stereognosis is
only an associated interpretation of all the impulses of the sense Tactile sensations appear of touch and not a subdivision of it.
to
Spiller
and
Mills,
may be
conducted by
sensations
common
sensory nerve
fibers.
Other common
seem
to require
some
pulses pursue a path entirely In giving the common sensory pressions of the muscular sense. the following classification will be adhered to, though tracings,
is still
lacking.
tactile
muscular and
joint surfaces,
and the
Spinal and cerebral (Fig. 107). II. Paths carrying impulses of the muscular and tactile sen-
ses, chiefly
from viscera
III.
Gracilis et
Cu-
neatus (Fig. 107). Impulses originating in the end-organs of the spinal nerves traverse the dendrites of the spinal ganglion neurones
(Cajal), the cell-bodies in the ganglia,
of the
same.
They
378
spinal nerves,
TRACING OF IMPULSES.
and ascend through the posterior column as far as the eighth thoracic segment and then through the fasciculus gracilis, or, entering above the eighth thoracic segment, they
ascend through the fasciculus cuneatus.
In either case they arrive
in one of the nuclei of the posterior column, namely, the nucleus
funiculi gracilis or the nucleus funiculi cuneati.
by a direct or by an indirect route. pulses may The direct route carries the impulses by way of the medial (1)
proceed either
fillet
pons and mid-brain, to the lateral nucleus of the from which they are conducted by the cortical fillet thalamus,
reticularis of
to
In their
last
stage the impulses run from the thalamus through the internal capsule and corona radiata to the posterior central gyrus in the
equatorial zone of the hemisphere. (2) Indirect Route. By that route impulses from the nucleus funiculi gracilis and nucleus funiculi cuneati run to the cortex
of the vermis cerebelli superior through the external arcuate fibers then on, through the brachium conjunctivum, to the red nucleus and thalamus. They traverse the restiform body of the same
;
side,
by way
by way
fillet
decussation of the medulla and the opposite restiform body to reach the vermis cerebelli superior. From the cerebellar cortex,
the impulses continue through cortical axones to the nucleus dentatus, whose axones conduct them to the red nucleus and
side. The greater number, therefore, tegmentum of the mid-brain. Their course from the red nucleus and thalamus is through the cortical fillet
to the cortex.
These impulses from the spinal nerves go to the upper twothirds of the posterior central gyrus, those from the lower extremity to the upper third and those from the arms to the middle
third (Spiller).
crossed fibers from the terminal nuclei of the trigeminal, the vestibular, the glossopharyngeal and the vagus nerves join the
379
Brachium conjunctivum
Pons and cerebellum
V nerve
body
IX nerve
Medulla
X nerve
Gowers
Tract
Spino-thalamic tract
Spinal nerves
Cord
Common sensory paths, pain, temperature Fig. 108. anterior cerebello-spinal and spino-thalamic tracts.
;
and
Posterior root-fibers connected with this path end in the center of the crescent and in the base of the anterior columna of both sides and the ascending fibers rise partly on the same and partly on the opposite side; the crossed fibers run through the white anterior commissure.
381
muscular and tactile by those cerebral nerves to their nuclei in the medulla and pons, are carried by the medial fillet to the lateral nucleus of the thalamus on the opposite side. The cortical fillet conducts them to the lower portion of the posterior central
to the thalamus, so
and run
sensations transmitted
gyms
H.
PATHS CARRYING IMPULSES OF THE MUSCULAR AND TACTILE SENSES, CHIEFLY FROM VISCERA (?).
(Posterior) Cerebello-spinal Tract (Direct CereThe column of cell-bodies, forming the nucleus bellar) (Fig. 107).
Through
dorsalis
(Clarki),
once to the (posterior) cerebello-spinal tract, through which they ascend along the dorso-lateral surface of the cord, along the
posterior surface and through the restiform body of the medulla to the cortex of the superior worm of the cerebellum. To a smaU extent they cross in the worm to the opposite side. These
impulses probably excite, within the cerebellar cortex, impulses of coordination and equilibrium which traverse the descending
anterior cerebello-spinal tract to the gray crescent of the cord, and are transmitted to the motor neurones of the anterior columna.
From
the cerebellar cortex the journey to the cerebrum is completed in four stages, as already described, namely: Nucleus dentatus,
opposite red nucleus, thalamus and some part of the cerebral cortex, probably the middle and inferior temporal gyri.
Probably the vagus nerve conducts impulses from the viscera to its terminal nucleus; but,
of similar nature
if
so,
their path
is
TEMPERATURE AND TACTILE IMPRESSIONS. SPINAL AND CEREBRAL. Through Spino-thalamic and Ascending Anterior CerePATHS CONVEYING
PAIN,
bello-spinal Tract (Fig. 108). In the spinal cord, medulla and pons these constitute one tract, commonly called Gowers's tract.
They
separate just below the isthmus, whence the spino-thalamic thalamus and the other turns back to the
cerebellum.
They appear
to
382
TRACING OF IMPULSES.
temperature impulses. These impulses enter the gray crescent A of the cord, on both sides, through the posterior nerve roots.
large
number
commissure; the
through
the
decussate via the posterior nerve roots in the gray rest decussate in the first stage of the ascending
spino-thalamic
cerebello-
thalamus and to
In the cord they
ascend along the lateral surface. They run dorsal to the olive in the lateral area of the medulla oblongata, and through the lateral
part of the formatio reticularis of the pons to the angle in Gowers's tract situated near the isthmus. From the angle, just below the
thalamus, and from that to the posterior central cortex. The common course of sensory impulses from the cerebellar to the
cerebral cortex
is,
and brachium conjunctivum to opposite red nucleus and thalamus. Having arrived in the thalamus, they proceed thence by
the cortical
fillet
on through the restiform body to the cerebellum by the tract from the lateral nucleus to the cerebellar cortex, thence to the
somaesthetic area as previously given.
108).
and trigeminal nerves those nuclei they .are conducted by axones which probably enter into the spino-thalamic tract, and, perhaps, into the ascending anterior cerebello-spinal tract,
certain fibers of the vagus, glossopharyngeal
to their terminal nuclei.
From
to the
now
What
special varieties of
common
383
sensation are conducted through these paths is unknown. Under certain conditions, perhaps, they may carry all varieties, (i) The antero-lateral fasciculus proprius and formatio reticularis contain
ascending axones which may convey sensory impulses from the gray matter of the cord, received from the posterior roots of the
spinal nerves,
in
common
ward
to the
is
The
(2)
thalamus
by way
of the cortical
Again, impulses
may
tis,
brachium pontis
by way
of
dinary course through the brachium conjunctivum to the red nucleus and thalamus.
Destruction of any of the above sensory paths causes diminution or loss of the especial variety of impulse
which
travels that
loss of
posterior produces muscular sensations and gives rise to ataxia. Interruption of Gowers's tract (spino-thalamic and ascending anterior
path.
Destruction
of
the
white
columns
while touch
is
not
much
2.
affected.
Special Sensations.
sight,
and
hearing to the
brain by the following nerves: The olfactory; the optic; the auditory; and the glossopharyngeal and intermediate nerves.
Olfactory Path
(Figs. 109
and
21).
inate in the upper third of the nasal mucous membrane. They run through the olfactory nerves to the second layer in the bulb,
cells.
where they are transferred to the dendrites of the mitral and brush By the axones of these cells they are carried backward
through the olfactory tract and striae to the cerebral hemisphere. The medial stria conducts them to the parolfactory area (Brocae),
the triangle, the gyrus subcallosus and anterior end of the gyrus cinguli, whence, through the cingulum, the fornix and uncinate
fasciculus they
may
384
hippocampi.
TRACING OF IMPULSES.
By
the intermediate olfactory stria the impulses and anterior perforated substance, and com-
same manner
as given above or
The
Central core of
ependymal
cells
Fibers to Ol.
tract.
5.
S.
granulosum
4. S. cellulare
3.
S. reticulare
2. S.
glomerulosurn
i.
S.
nervosum
Nasal mucous
membrane
Olfactory cellbodies
Fig. 109.
chiefly, if
not entirely,
in the rods
series of
Optic Path (Figs, no and in). Impulses of sight originate and cones of the retinae and traverse three or more
neurones to the terminal nuclei of the optic tracts; namely, rod and cone, the bipolar, and the ganglionar neurones. axones of the last form the optic nerves and the visual part
the
The
385
From
halves of both, impulses run through the correspondtract to the lateral geniculate body and the pulvinar of the ing thalamus; also to the superior quadrigeminal colliculus. The
from the
left
latter
reflexes.
From
the lateral
Fig.
Cells, s'
2'.
no.
The
(After Brubaker.)
Visual cells with their peripheral terminations, s. Rods. z. Cones, b. Bipolar cells, g. Ganglion cells from which arise the axones of the optic nerve.
side,
temporal halves, for the most part at least, remain on the same but a few may cross through the quadrigeminal colliculi
the temporal half of the macula lutea are conducted equally by both optic tracts. Hence destruction of one tract causes hemianopsia, preserving the vision in the corresponding half of each visual field, and also diminishes the acuteness of macular vision
in both eyes.
Auditory Paths. There are two auditory paths, cochlear and the vestibular. The former is concerned with hearing and
the latter with equilibrium.
25
386
TRACING OF IMPULSES.
1. The Cochlear Path (Figs. 89, 70 and 54). Impulses of hearing originate in the organ of Corti. They are transmitted by the rods and hair cells of Corti to the dendrites of the spiral gan-
Traversing the dendrites and cell-bodies of that ganglion, they enter the axones, which form the cochlear nerve, and run backward to the terminal nucleus of that nerve hi the medulla.
glion.
From
and dorsal
to the restiform
the opposite side through the medullary striae and trapezoid body, or they run medial to the restiform body and enter at once into
the trapezoid body. By either course they reach the lateral filand chiefly the opposite one. The lateral fillets conduct the let,
impulses to the inferior quadrigeminal colh'culi; the brachia inferiora to the medial geniculate bodies, and the temporo-thalamic
radiations to the third
and fourth
fifths of
and
Through
the lateral
by the
pass to
motor
and
sixth,
Vestibular Path.
is
The
path
and semi-
and
the path of space sense. Through the vestibular nerve the impulses reach the dorso-medial, the dorso-lateral and superior nucleus,
fourth ventricle.
pursue, from the terminal nuclei in the ventricular floor, either a direct or an indirect course to the cerebral
The
may
cortex.
(i)
fillet
By
and
387
colliculus
superor
corpora quad.
Fig.
in.
The
optic path.
(Original.}
389
impulses run to the cerebellum, by the indirect course, the descending root, and the external arcuate fibers in through the restiform body. They thus reach the cerebellar cortex. They
The
may
and then
the cerebellum the course of the impulses is, presumably, through the brachium conjunctivum to the red nucleus and thalamus of both sides and thence to the cortex.
continue upward.
From
Impulses, believed to be concerned with reflexes, run from the vestibular nuclei in the floor of the fourth ventricle, (a) to the opposite nuclei of
bundle; (b) to the quadrigeminal colliculi through the superior fillet; (c) to the cerebellum by way of the descending root and arcuate fibers, whence they reach the spinal nerves through the
descending anterior cerebello-spinal tract; (d) to the spinal cord and nerves through the axones of Deiters's nucleus; and, perhaps,
less directly
fibers
of Kolliker.
solitarii in the medulla and thence, probably, the opposite formatio reticularis and internal capsule through to the taste area in the gyrus fusiformis (Mills) or gyrus cinguli There are two paths from the tongue to the nucleus (Flechsig).
Those impulses from the base of the tongue and the palate run through the ninth nerve and those from the anterior two- thirds of the tongue through the chorda tympani and intermediate nerve to the medulla (A. F. Dixon, Keen and
of the solitary tract.
Spiller,
H. Gushing,
the
palate
etc.).
ating
in
may
Meckel's ganglion and the great superficial petrosal nerve to reach the geniculate ganglion on the facial; and, then, continue through the intermediate nerve to the solitary tract. All impulses arriving at this nucleus of the solitary tract probably complete their journey
in
site
First, through the formatio reticularis to the oppothalamus; and, second, through internal capsule to the cortex. Destruction of the olfactory conduction path on one side
:
two stages
causes anosmia on the same side; of the optic tract or radiation, atrophy and destruction in the corresponding halves of both retinae
3QO
TRACING OF IMPULSES.
above the pons, deafness
chiefly
on the
opposite side; and interruption of the gustatory path above the taste on the opposite (?) side.
REFLEX PATHS.
There is no visible limit to the number of reflex paths. Hence no attempt will be made to give them completely, but a few examples of various kinds will be given which may assist the student to trace others and be suggestive of their great multiplicity and
importance.
Under
sory and motor paths that have been traced are but the afferent and efferent limbs of reflex arcs.
(i)
fibers
which are
by
by the ascending
fibers of the
and (4) by afferent cerebral and efferent the two being associated by the anterior
longitudinal bundle, the ponto-spinal tracts, the fasciculi proprii, the spinal tract of the fifth nerve, the vestibulo : spinal tract, the
solitary tract, etc.
(i)
Spinal Reflexes
spinal reflexes,
cell-bodies
In the simplest the afferent fibers of the arc arborize about the
(Figs.
112 and
113).
ent
and
by one
neurones in the next grade of reflex arc. Among these are the skin and muscle reflexes, such as the plantar, the patellar, the
gluteal
of a part
reflexes, the involuntary withdrawing from a source of irritation, etc. Coordinating and equilibrating reflex impulses traverse much
longer arcs. They are composed of the spinal ganglion neurones, the external arcuate fibers (and probably the [posterior] cerebellospinal tract) in the afferent limb,
cerebello-spinal tract and the anterior root neurones in the efferent limb of the reflex arc.
REFLEX PATHS.
391
complicated spinal reflexes are those of defecation, micturition, parturition, vasomotor reflexes, cardio-accelerator reflexes, etc. The impulses traverse at least three neurones in these reflexes;
More
because
all efferent
organ supplied.
As an example,
trace
Defecation Reflex. The rectum is supplied by the third and fourth sacral nerves and by branches of the inferior mesenteric
the
and hypogastric plexuses. Irritation of the sensory endings in mucous membrane is caused, normally, by the presence of
feces.
center in the
lumbar enlargement
by
Fig. 112.
i.
Diagram
2.
(After Brubaker.)
cell.
Sentient surface.
Afferent nerve.
5.
Emissive or motor
4. Efferent
nerve-
Muscle.
way
the ganglionated cord, and the rami communicantes to the lumbar which they reach the center the defecation center the impulses pursue (a) They descend through the third and fourth sacral nerves and cause inhibition in the circular fibers of the
From
two courses:
rectum and contraction of the longitudinal muscle, (b) This action is immediately followed by impulses which pursue the
sympathetic course, through the anterior roots of the lumbar nerves, the rami communicantes, the ganglionated cord, and the inferior mesenteric and hypogastric plexuses, to the rectum.
They
cause,
in succession
39 2
TRACING OF IMPULSES.
it
thus open a way for the passage of fecal matter; and, then, force through the opening unless prevented by the voluntary contraction of the external sphincter.
(2)
Cerebral Reflexes.
The
such as spasm of the muscles of mastication caused by a bad tooth, in which both limbs of -the arc are formed by the trigeminal
nerve.
cause.
Again, the facial expression of pain due to the same In this the impulses traverse the trigeminal nerve and
Fig. 113.
A more complicated
(Brubaker after
Diagramjshowing the relation of the third neurone a, to the afferent neurone efferent neurones c, c, c.
and
to the
by the collaterals of its root-fibers reach the nucleus of the facial. Through the facial they cause contraction of certain muscles of Facial spasm in tic douloureux is due to the same expression.
reflex.
The
involuntary expansion of the nostrils upon the detecis due to an The conolfactory-facial reflex.
nection of the terminal nucleus and cortical center of the olfactory nerve with the genetic nucleus of the facial nerve is very much
REFLEX PATHS.
in doubt;
it
393
may be
by the
fornix, the
(2)
fibers,
by
the fornix, medullary stria, fasciculus retroflexus and descending fibers of the interpeduncular ganglion, and, (3) by the fornix and the pedunculo-mamillary bundles. Squinting, due to bright light, produced by an arc composed of the visual path, the corticifugal
is
part of the occipito-thalamic radiation, the anterior longitudinal bundle and the facial nerve. Substitute the oculomotor nerve
for the facial
the
same
conditions.
Salivary reflexes, in which the sight of a fine dinner or the smell of it causes the flow of saliva coughing, sneezing, vomiting reflexes and deglutition reflexes are complicated, but, knowing the nerve
;
supply of the parts involved, the student should try to trace the
impulses.
(3)
Spino-cerebral Reflexes.
cord through the afferent fibers of its nerves are transmitted by the medial longitudinal bundle, the fasciculi proprii and formatio
motor cerebral nerves. Thus is brought about the movement of the head and eyes toward the source of impulse, a change of facial expression to agree with the painful
reticularis to the nuclei of
etc.
Cerebro-spinal Reflexes.
Of
many.
Let
us notice three.
Respiratory Reflex. Any obstruction or irritation in the larynx or trachea sends an impulse through the vagus nerve to its sensory nucleus and, through its descending branches, in the solitary
tract, to the
spasm
Equilibrium Reflex (Vestibulo-spinal Reflex). The simplest arc of equilibrium between the eighth cerebral nerve and the spinal nerves is formed by the neurones of the vestibular ganglia
(Scarpa's), the vestibulo-spinal part of the descending anterior
cerebello-spinal
columna
of
and the motor neurones of the anterior the spinal cord. A more complicated arc includes
tract
394
fibers
TRACING OF IMPULSES.
from the nucleus
funiculi cuneafi
cerebello-spinal tract.
Pupillary Reflexes.
spinal group
of reflexes.
enlargement of the spinal cord. It receives optic impulses through the anterior longitudinal bundle from the corpora quadrigemina. The superior quadrigeminal colliculi receive those impulses by
two routes:
of the optic
First, directly,
tract,
through the
By
the visual area of the occipital lobe, by way of the intrinsic retinal neurones and the optic nerves, tracts and radiation, are returned
to
the
lateral
Thence,
geniculate and superior quadrigeminal bodies. reaching the cilio-spinal center through the anterior
longitudinal bundle, the impulses take the following course: They leave the spinal cord through the anterior roots of the upper thoracic nerves and run, in succession, through the rami communicantes, the cervical cord of the sympathetic, the cavernous plexus, the ciliary ganglion and the short ciliary nerves to the radiating fibers of the iris, causing dilatation of the pupil.
For pupillary
superior
traversing
constriction, the
quadrigeminal
the
anterior
colliculus
longitudinal bundle only through the dorsal tegmental decussation (Meynerti). Then, through the third and short ciliary nerves, they reach the sphincter pupillse
muscle.
CHAPTER
VIII.
line, in
Very soon after conception there appears along the median the back of the embryo, a plate of epiblast, called the
medullary plate. That plate soon presents two longitudinal elevations, the medullary ridges, separated by a median furrow, the neural groove (Fig. 16). At first the medullary plate is formed of
a single layer of columnar cells; but those cells undergo rapid mitosis, as the medullary ridges are developed, and arrange themselves in several layers
(Fig.
114).
The
rapid growth, roofs over and closes hi the neural groove and produces the neural tube (Fig. 16). The tube is first formed in the
cervical region,
in
of the medullary ridges extends the approximation of the medullary ridges By a slight paramedian crest is produced on either side,, called the neural crest. The neural tube and the neural crest are nearly
both 'directions.
is
open
median aperture
of
Magendie.
Both ends
closing
first.
neural tube
is
open for a short time, the anterior later grows around the developed into the meninges and the surrounding
bones, which inclose the cerebro-spinal axis. The Neural Tube (Figs. 16 and 114).
It is well
formed
at
the fifteenth day. The cephalic end of the tube is much larger than the caudal end, and presents at this time two constrictions
that separate the primary brain vesicles from one another the anterior, the middle, and the posterior. Behind the posterior
395
39 6
primary uniform
plan of
forms the spinal cord, in development may be observed in its simplest form.
size; that part
IV.
Sections showing stages in the conversion of the medullary groove into Fig. 114. the neural canal. From the tail end of an embryo of the cat. (Gordinier after E. A. Schafer.) ep,me,hy. Epiblast, mesoblast, and hypoblast. nt.g. Medullary groove, n.c. (in IV). Neural canal, ch. Notochord. cae. Celom. am. Tail-fold of the amnion.
make up
In the very early stages of development two varieties of cells the neural tube (Fig. 115). Certain very long cells,
397
are the ependymal cells, which form a supporting framework for the more important elements. They reach from the lumen of the tube to its peripheral surface
They
peripheral parts of the ependymal cells, by irregular branching and vacuolation, form a protoplasmic network destitute of nuclei, called the marginal velum. This marginal
The
Fig.
115.
Two
histologic layers in the embryonic spinal cord, (McMurrich after His.) long.
their bases.
embryo
4.25
mm.
among
The
ependymal
cells
and,
among
possessing large nuclei and small protoplasmic bodies (Fig. 115). The destination of the latter cells cannot yet be determined, so
they are
named
the indifferent
cells
(Fig.
116).
In the nuclei
all
may be
seen.
In a somewhat later stage of development the indifferent cells wander outward toward the marginal velum and arrange themselves into a definite layer,
known
and
tle
127).
manand the ependymal layer formed by the bases of the epenlayer, dymal cells. As the indifferent cells multiply in the mantle layer they become specialized into two types, the neuroblasts and the
in a cross section of the neural tube
Fig. 116.
The
c:ircles, indifferent cells; circles with dots, neuroglia cells; shaded cells, germinal cells; circles with cross, germinal cells in mitosis; black cells, nerve-cells.
spongioUasts.
The neuroblasts
They
are the
embryonic neurones. Each neuroblast throws out a protoplasmic process which develops into an axone and grows outward into the
marginal velum. Within the marginal velum the axone either runs some distance and returns to the mantle layer, or it passes
outward through the marginal velum and becomes an efferent or motor root-fiber. Collaterals are given off from the axones, the
formed and the medullary sheath is laid down as development proceeds and function begins. While these events
end-tufts are
399
are occurring in the efferent part of the neuroblast, other processes, called dendrites, are given off by the cell-body. They belong to
cell.
They branch
rule,
richly close to
The spongioblasts
They are primitive neuroglia cells (Fig. The spongioblasts develop many very slender and richly 53). branched processes of ray-like or arborescent form, which by their
interlacement constitute a delicate supporting reticulum. The neurones are held in the meshes of this reticulum, and both the
cells
are
still
While the neurones and neuroglia there occurs in each lateral wall of the neural forming,
Fig. 117.
embryo
Transverse section of the cervical part of the spinal cord of a of six weeks. (Gordinier after Kolliker, from Quain.)
human
c. Central canal, e. (Superiorly) The original place of closure of e. Its epithelial lining, the canal, a. The white substance of the anterior columns, g. Gray substance of anterior columna. p. Posterior column, ar. Anterior roots, pr. Posterior roots.
longitudinal groove
canal a longitudinal evagination. This evagination produces a on the ventricular surface of the neural tube,
which divides
zone
is
it
into a ventral
and a dorsal zone. The ventral hence from it issue the out127 and 128). It is composed of a median
on
The
by
dorsal zone
is
made
of
up two
400
the
median roof-plate. The dorsal zone is afferent in function and in connection with it the sensory nerves terminate. The roof-plate and the floor- plate take little or no part in forming neurones; they lack the mantle layer,
of neuroglia
and ependymal
cells.
(Figs. 16
and
The
cephalic por-
Those ganglia are called the vagus, glossopharyngeal, geniculate, auditory and semilunar (Gasseri). Later, the auditory ganglia lie between the geniculate and glosThese five ganglia give origin to the sopharyngeal ganglia. of the vagus, glossopharyngeal, intermediate and sensory parts In all the gantrigeminal nerves; and to all of the acustic nerve.
the brain.
glia except the auditory, the cells
dition is produced by the growth of the cell-body toward the surface of the ganglion and the shifting of both processes to the same side of the cell-body, together with the elongation of the common
The single processes of the unipolar neupoint of attachment. rones immediately divide, T-like, into peripheral and central fibers,
which in appearance are axones.
The
sensory part of the respective nerves and conduct impulses toward the cell-body, hence they may be considered dendrites (Cajal)
the central fibers, the axones proper, form that part of the nerve which extends from the ganglion into the dorsal zone of the em-
bryonic brain.
divide, T-like,
All
upon
the central axones of the several ganglia entering the brain, and collaterals rise from the
These axones
fibers, ter-
and
pective nerves; but certain of them, the excito-reflex minate hi nuclei of motor nerves.
The development
and
spinal,
of peripheral common sensory neurones, both cerebral has been observed in certain lower vertebrates. In the amblystoma
i. The elongation of the the neural crest to a spindle-form. 2. The projection of a slender central process, the axone, which grows into the neural tube. thick 3.
THE BRAIN.
irregular
4OI
dendrite, is thrown out from the peripheral end of the becomes smooth and fibrillar in character. This bipolar condition persists in very low forms, such as the amphyoxus and cyclostomes. The greater num4. In bony fishes some of the neurones become unipolar. ber become unipolar in higher vertebrates (/. B. Johnston's Nervous System
process,^
cell; later it
of Vertebrates).
(2)
The
of
pairs
spinal portion of the neural crest forms the thirty-one spinal ganglia situated on the posterior nerve roots;
sympathetic system. The sympathetic ganglia wander widely. In them the epiblastic cells develop into multipolar neurones,
the nonmedullated processes of which constitute the larger
num-
ber of gray fibers in the sympathetic system. The cells of the spinal ganglia form unipolar neurones, like those of the vagus,
glossopharyngeal, geniculate and semilunar ganglia. Like them, also, the single processes divide, T-like, the peripheral arms of the T-branches forming the sensory part of each spinal nerve
and the
nerves.
central
arms
The
The
descending
-fibers,
and end in the gray matter of the posterior the crescent and the anterior columna, the columna, the center of long fibers forming the postero-medial and postero-lateral descending radicular tracts; the ascending axones and collaterals terminate in the gray substance of the spinal cord of
its
posterior columns, namely, the nucleus funiculi gracilis and nucleus funiculi cuneati of the medulla. The long ascend-
ing fibers form the postero-medial tract (Goll's) and, the posterolateral tract (Burdach's); and, the shorter ones, those reaching
make up
The
greater
number
THE BRAIN.
The primary
fourth
week a
constriction
brain vesicles grow rapidly. By the end of the is visible in the anterior primary vesicle
26
402
and another
two and making in all five secondary brain vesicles, which freely communicate with one another and are numbered from before
backward.
They
are:
1.
Telencephalon.
2.
Diencephalon.
3.
Mesencephalon.
Metencephalon.
Myelencephalon.
4.
5.
form the brain, their cavities becoming the venand 17). The neuroblasts of the mantle the neurones, whose cell-bodies and dendrites are layer produce found in the cortex and ganglia, and whose medullated axones
vesicles
tricles
These
Flexures
is
The cephalic portion of the neural tube (Fig. 118). the seat of three flexures, two ventral and one dorsal, (i) The
mesencephalic flexure (ventral) begins very early and amounts to nearly 180 degrees by the twenty-eighth day. It bends ventrally
the diencephalon until
it
Thus
the inter-brain and pons are approximated and the mid-brain almost concealed. (2) The cervical -flexure is also a ventral one.
It is located at the junction of the fifth vesicle
cord,
and corresponds
to the
of the embryo. This flexion begins about the twenty-first day. the end of the fourth week, it is completed and amounts to By
90 degrees.
(3)
The
dorsal flexure
It
is
same time
(fourth week).
brain vesicles, and is often called the metencephalic -flexure. It reaches 180 degrees by the eighth week, when the dorsal part of the metencephalon (the cerebellum) rests upon the medulla
oblongata.
The
convexity
flexure.
synonym, pontine
permanent.
formed by the pons, hence the The cervical and metencephalic flexis is
THE BRAIN.
403
TABLE
IV.
and
120).
Primary
Vesicles.
Secondary Vesicles.
Telencephalon, Lateral ventricle and
Derivatives.
Anterior
the Pros-
Aula of third
tricle.
ven-
or
Diencephalon, and
Third
y Cerebrum.
ventricle, except
the aula.
Middle
the
Mes\
encephalon, Mid-brain.
or
l
Mesencephalon, and
aqueductus cerebri.
Posterior
lon.
the
(
J
I
Isthmus rhombencephali
!
Rhombencepha-
Rhombencephalon and
Fourth
ventricle.
Metencephalon
Myelencephalon.
TABLE
V.
Neo pallium
Rhinencephalon
(.
Corpus striatum.
Telencephalon
(End-brain)
(
Chiasma opticum
Thalamus
(
Thalamencephalon
Diencephalon
(.Inter-brain)
Metathalamus
.
Corpora
Geniculata
Epiphysis, and
(
f
Epithalamus
f
j
Corpora mammillaria.
404
Pedunculi cerebri.
4
(.
Tegmenta.
Corpora quadrigeminaColliculi superiores Colliculi inferiores
Mesencephalon
f
(Mid-brain)
Brachia
Brachia superiora Brachia inferiora.
Isthmus
cephali
are)
(in
Velum medullare
Cortex
superius.
Metencephalon
Corpus medullare
Cerebellum,
Ganglia
(Pedunculi cerebelli)
Vela medullaria.
Pons
Myelencephalon
-I
Varolii.
Medulla oblongatz
In the following resume of the formation of the central nervous system, I have drawn so largely from Prof. McMurrich's "Devel-
opment
grateful
of the
point, to
make
him.
TELENCEPHALON.
TABLE
VI.
DERIVATIVES OF TELENCEPHALON.
(Modified from McMurrich's Embryology.)
In Median Structures.
(
In Hemisphere.
\
(.
Roof-plate
(
Lamina
1T DorsalLamma
.
Lamina
_
Neopallium
Rhinencephalon
Optic
vesicle.
Corpus striatum.
THE BRAIN.
Anterior part of hypothal-
405
Ventral Zone
Infundibulum.
(
-.
Falx cerebri
Part of chorioid
ventricle.
t
.
tela of third
-<
Mesoblast
Meninges.
/
v.
/
v.
The telencephalon is at first a single vesicle forming the fore part of the anterior primary vesicle, and for a time is open in front at the neuropore (Fig. 118, VI). The median portion of its anterior wall,
it is
laterally,
the seat of rapid growth. Optic Vesicle. Almost before the telencephalon
differen-
from diencephalon, a club-shaped diverticulum, called the optic vesicle, is thrown out from the ventral part of its dorsal zone. It grows outward and forward, separates from the telentiated
cephalon and, becoming indented, forms the optic cup, from which
the retina
is is developed. indicated in the adult
The
by a slight
point of origin of the optic vesicle pit, termed the optic recess.
The
optic recess
is
and the
fornicis at the anterior end of the sulcus hypothalamicus (Monroi). This sulcus, in the anterior primary vesicle, separates the ventral from the dorsal zone.
columna
Hemisphere
after the
of
Cerebrum
(Figs. 17,
Soon on
either side in the dorsal zone of the telencephalon. That bulging produces a hollow diverticulum, whose cavity is the primitive lateral ventricle and whose walls form the substance of the hemisIts phere. The outgrowth is called the hemisphere vesicle. constricted stalk contains the primitive interventricular foramen
The
vesicles
at
separated from one another by mesoblast which forms the falx. Later, growth occurs in succession, upward, backward and
downward,
until
every other part of the brain. The hemisphere vesicle is formed almost wholly by the dorsal lamina of the telencephalon which
will
be considered further
406
on.
It
narrow prolongation
of the roof-plate.
This prolongation of the roof-plate undergoes very little development. At first it is placed in the roof of the hemisphere vesicle; but the upward growth of the hemisphere shifts it to the medial
surface and, later, the
carries
of the
hemisphere roof-plate through a horse-shoe curve down toward the pole of the temporal lobe. No thickening occurs in this prolongation. Becoming indented longitudinally it forms
the
the floor of the chorioidal
-fissure
in
which
is
developed, from
JY2
Fig.
1 1 8.
Median
month.
(McMurrich
I.
after His.)
encephali.
i,
Myelencephalon. II. Metencephalon: i, Pons, 2, Cerebellum. ITJ. Isthmus rhombIV. Mesencephalon: i, Pedunculi, 2, Corpora quadrigemina. V. Diencephalon: Pars mammillaris hypothalami, 2, Thalamus, 3, Epithalamus. VI. Telencephalon: i. Pars optica hypothalami, 2, Corpus striatum, 3, Khinencephalon, 4, Neopallium.
Rhinencephalon
and
120).
In the
fifth
week
a hollow diverticulum grows out from the antero-inferior wall of the hemisphere vesicle and, forms a prominent lobe. It preserves a lobular form in the horse, and in some other animals; but in man it soon becomes constricted by the sulcus parolfactorius
posterior into
an
anterior
tricular cavity.
The
and a posterior part, and loses its venanterior part develops the olfactory bulb,
THE BRAIN.
407
Transverse sections through the fore-brain of a four and one-half weeks embryo. (Gordinier and Quain after His.) A. Through the lower anterior part of the fore-brain. S. Falx. Sf. Fold of roof passing below falx toward the third ventricle. Bf. Fold forming the fissura hippocampi. v.Rl.,k.Rl. Anterior and posterior parts of olfactory lobe. Cs. Corpus striatum. O.W. Groove continuous with optic stalk. P.s. Hypothalamus. T.c. Tuber cinereum. B. Section a little farther back. Sf is replaced by a less prominent but broader fold of the roof, Ad, which subsequently receives the chorioid vessels, and is, therefore, the chorioid fold. Hs. Hemisphere vesicle. Th. Thalamus. S.M. Sulcus hypothalamicus (Monroi), below and behind the thalamus. C. Still farther back. Ad. Chorioid fold here projecting into lateral ventricles, but still free from mesoblast and blood-vessels. Ma. Mammillary tubercle. The other lettering as
Fig.
1 19.
before.
THE BRAIN.
tract,
409
triangle, and the pardlfactory area (Brocae), the posterior portion forms the anterior perforated substance, etc. Primary Fissures (Figs. 121 and 122). The vesicle walls are
uniform thinness up to the second month. During their rapid growth in the second and third months they become thrown into
of
folds
The
cause
cranium.
The
They
is, perhaps, the resistance of the slower growing primary fissures are best developed in the third
month.
is
concentric with
Fig. 120.
Diagrammatic
(Morris's
Anatomy
a. Corpora quadrigemina. b. Mid-brain, c. Pineal body. d. Cerebellum (hind-brain). Medulla oblongata (after-brain), f. Pons Varolii (hind -brain), g. Lateral ventricle. h. Cerebral hemisphere, i. Corpus striatum. j. Olfactory diverticulum. k. Pedunculi cerebri. 1. Thalamus, m. Inter-brain, n. Hypophysis, o. Interventicular foramen. 4. Fourth ventricle, s. Aqueduct of cerebrum. 3. Third ventricle.
e.
it,
lateralis cerebri
(Sylvii).
The hippocampal
fissure (Fig. 121) begins near the frontal pole on the medial surface of the hemisphere vesicle and extends backward, downward
It divides the lastly, forward toward the temporal pole. medial surface into a broad marginal gyrus and the dentate gyrus, the latter is between it and the chorioidal fissure. In its whole
and,
course
it is
and
121).
The
is by called the hippocampus. two The hippocampal fissure gives off branches at its most posterior part, which represent the occipitoBoth branches proparielal and the calcarine fissures (Fig. 121).
410
sists,
entirely disappears.
The primary occipito-parietal fissure The whole superior part of the hippocam-
pal fissure
is
losal fissure.
represented in the adult, as to position, by the calThe lateral cerebral fossa (Fig. 122) is a deep bay
hemisphere
vesicle.
It is
due
to the
relatively limited growth of the corpus striatum in comparison with the more rapid and greater growth of the surrounding parts.
The
thickened floor of the fossa cerebri lateralis develops, interforms the insula (Reili).
The lateral fossa and the four fissures mentioned above are well formed by the third month, but the lateral cerebral fossa is not converted into a fissure until the end of the fifth month. Even
is
then only the posterior ramus of the fissura cerebri lateralis (Sylvii) formed, and this is brought about by the meeting of the temporal
fronto-parietal parts of the operculum.
after birth that the
It is
and the
first
during the
year
and
frontal parts of the operculum produces the anterior horizontal ramus and the anterior vertical ramus of this fissure.
(Fig.
123).
The
they
About the middle of the fifth month the first secondary sulcus makes its appearance. It is the sulcus cinguli of the medial surface, which separates the superior frontal gyrus and paracentral lobule from the gyrus cinguli
of the adult brain (Fig. 27). pieces explains its In the sixth month the subparietal
present.
There
of the fifth
and occipito-parietal sulci are no sign of the latter sulcus in the early part month and the permanent sulcus does not produce a
is
ventricular eminence, hence the occipito-parietal indentation is a sulcus and not a fissure (see Cunningham Memoirs). Early in
the sixth
month
as,
formed, such
the most important of the remaining sulci are the central sulcus (Rolandi), the precentral, the
23 and 123). The only secondary fissure, the collateral fissure appears in the same month. It is developed on the tentorial sur-
THE BRAIN.
face of the hemisphere, at
terior
first,
411
an anmiddle
and a
is
posterior temporal
and an
part
a total
sometimes.
When
parallel
both are
with
total they
hippocampus and
name, the eminentia collateralis. There are many tertiary sulci which do not appear birth, some of them one or two years after birth.
hence
its
until
near
Medial sagittal section through the brain of an embryo of three months, Fig. 121. showing the primitive fissures on the medial surface of the cerebral hemisphere.
(McMurrich
c.
after Mihalkovicz.}
ca.
Calcarine fissure,
Anterior commissure, cc. Corpus callosum. cf. Chorioidal fisForamen interventriculare (Monroi). h. Hippocampal fissure.
of this fissure,
which
is
belongs wholly to the cerebrum; the posterior part separates cerebrum from cerebellum. The anterior part of the transverse
fissure is
bral hemispheres over the free dorsal surface of the diencephalon, the inter-brain. It is continuous laterally with the chorioidal
fissure in
tela of the
third ventricle.
As the
gyri of the
cerebrum
412
and
are acquiring their adult forms and producing the various fissures sulci, the walls of the hemisphere vesicle thicken rapidly and
give rise to the corpus striatum, the cortex and the medullary substance; but the order of development is not yet understood.
The development
of the gray
and the
the cavity of the hemisphere vesicle to the size of the lateral ventricle; and the cornua of the ventricle are produced by the forward, back-
ward, and downward growth of the vesicle in the successive formative stages. Little is known of the time at which the cortical
neurones are formed; but it would seem that they continue to undergo medullation and to become functional up to a late period
of
and the investigations of Kaes support this inference. Fornix. A ridge appears, about the fourth month, on the medial
life,
That
foramen.
It
which
is
lateral wall
It
and
floor of the
diencephalon
mammillare.
forms a
The union
of the
The body
is
extended by the
and by
the crossing over of certain fibers from one crus fornicis to the other, which results in the formation of the commissura hippo-
campi.
Another ridge, a slight one, appears about the same time on the opposite lip of the chorioidal fissure. It represents the stria terminalis. This latter ridge and the fornix ridge bound the roofplate as represented in the hemisphere.
undergoes no development. It forms the chorioid epithelial lamina of the lateral ventricle, which loosely joins the fornix and
the stria terminalis, being folded over the chorioid plexus of that
ventricle (Fig. 119).
Internal Capsule. It is formed largely along the line of fusion between the medial surface of the hemisphere vesicle and the
lateral surface of the diencephalon.
grow downward through the corpus striatum from the cerebral cortex, and
Motor
fibers
THE BRAIN.
413
sensory fibers ascend through the striated body to the cortex from the thalamus and other ganglia. All these motor and sensory
fibers together constitute the internal capsule. Its bell-shape in the hemisphere is due to the rotary growth of the hemisphere, upward, backward, downward and forward, around the corpus
Anterior Commissure
at the fourth
teriorly
(Fig.
121).
The lamina
terminalis
week
is
a thin median plate bounding the aula anvesicles together just in front
Its ven-
Fig. 122.
The
fossa lateralis cerebri, in embryonic brain of the fourth month. (After McMurrich.)
c.
Cerebellum,
p.
Pons.
5.
Fossa
lateralis cerebri.
with the upward and backward growth of the hemisphere, it is extended in a crescent, dorsal to the interventricular foramen,
as far
this
backward as the splenium of the corpus callosum. Within thickened crescentic part of the lamina terminalis are formed the anterior commissure, the body of the fornix, the corpus callo-
sum and
first
formed.
tion
the septum pellucidum. The anterior commissure is the ventral angle of the thickened porThrough grow, transversely, the commissural and the decussating
fibers of the
ing the occipito-temporal cortices. These fibers make up the anterior commissure. The fornix fibers grow for the most part
414
longitudinally through the zone next the ventral surface of the crescentic lamina and, on reaching the anterior commissure, bend
backward
a certain
of the
and
third ventricle;
number
lamina terminalis and produce the commissura hippocampi. The corpus callosum is formed in the dorsal zone of the crescentic
lamina terminalis.
(Fig. 121) is
produced by
fibers that
to the other
Fig. 123.
Permanent
fissures
as seen in a seven
is.
and sulci on the convex surface of the cerebrum months embryo. (McMurrich after Cunningham.}
PCS.
t
l
.
Superior frontal fissure, ip. Interparietal. IR. Island (Reili). pci. Inferior pre-central. R. Central (Rolandi). S. Lateral (Sylvii). Superior pre-central. ptc. Post-central. First temporal.
The
corpus
callosum
are
truncus, splenium growth in successive stages of cerebral development. The fibers of the corpus callosum pierce the dentate gyrus of the hemisphere
vesicle.
formed in regular order rostrum, genu, and this order coincides with the lines of
of the
In so doing, they completely obliterate the superior part hippocampus and leave but rudiments of the dentate gyrus,
and peduncle
of
DIENCEPHALON.
cinerea
415
tary remains.
That part of the lamina terminalis included between the corpus callosum and the body of the fomix persists as the septum pellucidum. It develops a lymph
Septum Pellucidum.
is
which
fifth ventricle
(cavum
septi
to
be translucent.
The above description, giving the origin of the septum pellucidum, corpus callosum and anterior commissure from lamina terminalis, is the one commonly accepted; but it is not their origin in the rabbit (Marshall) or in the and there is still room for doubt concerning their origin in the human brain. It is possible that all or a part of them are developed in a
rat (Zuckerkandl),
in
lower animals.
The
infundibulum are true derivatives of these telencephalic zones, but the optic chiasma is not. It is produced by the ingrowth of fibers from the retinae and the medial geniculate bodies.
DIENCEPHALON.
It
is the posterior division of the anterior primary vesicle. forms the inter-brain and contains the greater part of the third ventricle (Figs. 17, 118, V, and 119). Its lateral walls pre-
This
sent
on each
thalamicus (Monroi) (Fig. 119, B and C) which separates the ventral zone from the dorsal zone (?).
Roof epithelium of
Corpus pmeale
third ventricle.
Roof-plate
I i
(epiphysis).
,
Dorsal Lamina
Thalamus
.
{
I
Ventral Zone
(
I
Corpus mammillare
A small
.
Surroundmg Mesoblast
tncle.
416
The roof-plate
120).
Its
and becomes
118, V3,
and
major portion forms the epithelium covering the third Posteriorly, fibers grow through it from opposite sides and form a transverse white band, the posterior commissure. Implate appears
body
The pineal the primitive pineal body. soon becomes solid and is joined to the diencephalon by a
which
is
Fig. 124.
Dorsal view of an embryonic brain, the roof of the lateral ventricles having been cut away. Embryo of 12.6 mm. (McMurrich after His.)
superius. eg. Corpus geniculatum laterale. Cp. Chorioid'plexus collateral Cqa. Colliculus superior of corpora quadrigemina. h. Hippocampus, hf. Hipfissure. Ot. Thalamus. p. Pineal body. rp. Roof -plate.
b.
Brachium
ventricle.
pocampus and
is
ventricular cavity, called the pineal recess (Fig. 120). Anterior to the pineal body, two longitudinal folds of the roof-plate dip down into the ventricular cavity. These are followed by two
MESENCEPHALON.
like
417
projecting folds from the inferior lamina of the chorioid tela, which constitute the chorioid plexuses of the third
ventricle.
downward
and
upon
into a
narrow
Externally, the dorsal lamina fuses with the cerebral hemisphere, as a result of the formation of the internal cap-
median
slit.
and the fusion at one point of the medial surfaces of the two dorsal laminae gives rise to the massaintermedia (middle commissure). The development of the thickened dorsal lamina of the diencephasule;
its
floor-plate of the diencephalon conmammillaris hypothalami, which embraces some gray and white matter beneath the thalamus, and the corpora mammillaria (Fig. 118, VI). A single oval eminence situated
stitute the pars
in the
third
represents the mammillary bodies month; but, during the third month, that eminence
line,
median
up
is
to the
divided
into the
adult.
The
gray substance immediately in front of the corpora mammillaria also belongs to this region, hence a part of the tuber cinereum is
included
among
MESENCEPHALON.
This
is
the
mid-brain
is
and
120).
It is the
vesicles.
The elbow
it;
of the mesencephalic
formed by
and that
flexure almost
another beneath
it.
walls thicken greatly. As a result of the thickening, its cavity is reduced to a slender canal, the cerebral aqueduct (Fig. 120).
TABLE
f
VIII.
DERIVATIVES OF MESENCEPHALON.
Lamina quadrigemina
Colliculi
Dorsal Zones
~\
Brachia
,.
..
(in part)
[_
Red
nuclei.
27
41 8
fifth
the development of the mid-brain. The origin of the derivatives tabulated above is largely inferred from their position and function. The formation of the corpora
Little
is
known about
In the
dorsal part of the mesencephalon, an elongated eminence on either side of median line is present at the beginning of the third month.
Those two eminences resemble the corpora bigemina of birds, Two months later a transverse groove divides fishes and reptiles. each eminence into the superior colliculus and the inferior colliculus. By the growth of fibers through the marginal velum between the colliculi and the corpora geniculata the brachium superius and the brachium injerius are produced (Fig. 43). The bases
from the
pedunculi are produced, likewise, by the down-growth of fibers fore-brain, which traverse the marginal velum in the
METENCEPHALON.
The metencephalon is the fourth of the secondary brain vesicles. From it are derived the isthmus, the cerebellum and the pons
(Figs. 17, 118, III, and IV, and 120). Its dorsal wall, which forms the cerebellum, presents a transverse indentation, the metencephalic angle and, as a result of that angle the opposite wall is bulged
;
forward.
The
The
ventricle.
Superiorly, this cavity contracts to the size of the cerebral aqueduct; it expands inferiorly and is broadest at the junc-
The
ventricular
surface presents on each lateral wall a longitudinal furrow which divides the metencephalon into a ventral and a dorsal zone. In
is
indicated
by the sulcus
METENCEPHALON.
419
limitans of the rhomboid fossa, containing the }ovea superior and the locus caruleus, hence, the pons comprises the greater portion
of the metencephalon, including the entire ventral zone part of the dorsal zone.
and a
TABLE
IX.
(McMurrich, modified).
Isthmus
{
DERIVATIVES OF METENCEPHALON
Metencephalon Proper
Roof -plate
j
)
Inferior
velum
Cerebellar vermis
trochlear
f
|
Lobes of cerebellum
Terminal
nuclei, of sensory
fifth
Dorsal Zones
nerves (part of
and
i
Brachium conjunctivum of
cerebellum grows through
it.
?)
Nucleus pontis
Genetic
nerves
nuclei
(fifth,
?)
of
motor
sev-
Inferior
part
of
of
tegmen-
sixth,
tum and
\
substantia
Ventral Zones
enth)
~{
nigra.
medius of the
re-
ticular formation
Floor-plate
Median raphe
Median raphe.
Cerebellum.
It
down
cephalic flexure (Fig. 118, between II and I). Above the plica chorioidea the dorsal laminae of the metencephalon thicken
rapidly and form a prominent transverse ridge on either side of the median line. The two thickenings are partially separated
line
by a deep
ventricular sulcus
At
But cerebelli, the ridges represent the hemispheres. very soon, cells from the dorsal laminae invade and thicken the
vermis
42O
The
cerebellum
now represented by one continuous transverse ridge arching over the fourth ventricle. It develops slowly in comparison with
the cerebrum.
The
median portion
month.
more rapidly than the Thus the hemispheres The flocculus is the first
and
it
of the hemisphere
of the
human
lobe, the superior semilunar and the inferior semilunar lobules, are not fully formed until near birth. The jolium vermis of the worm is developed after birth (Figs. 76 and 80).
The
worm appear
in the
month; with two exceptions, those of the hemisphere develop later. The lateral part of the postnodular sulcus is first manifest. In the second month it cuts off a strip of the cerebellar
ridge, just
plica,
which
is
culus.
median
line
posterior
boundary
of the nodule.
The
next sul-
cus to develop is the predeclivil sulcus (Fig. 79). It cuts very deeply into the vermis between the culmen and the declive. From
it extends into the hemispheres, where it separates the anterior and the posterior parts of the quadrangular lobe. The prepyramidal and the postpyramidal sulci are formed near
the vermis
month
(Fig. 79).
behind
the quadrangular lobule (Fig. 76). Soon it becomes continuous It is the vermis with that of the opposite hemisphere. through the beginning of the fifth month before the prepyramidal sulcus
is
lobule (Fig. 80). At about the same time the lateral extension of the postpyramidal sulcus establishes the posterior boundary of the biventral lobule. According to O. C. Bradley's study of the
rabbit's cerebellum, the postcentral sulcus
and the
precentral sul-
cus aj>pear at about the same time as the postdeclivil. The last important sulcus to appear [is the horizontal sulcus (Figs, 76
METENCEPHALON.
and
80).
It is
421
is
produced
almost wholly by the enormous growth of the superior and the inferior semilunar lobules which are so characteristic of the human
cerebellum (Cunningham).
cerebellar lobules are subdivided into gyri by intralobular which develop in the later months of pregnancy and the early months of extrauterine life. There are two, called the midgrasulci
The
and postgracile sulci, which Bradley says are found only in man and the anthropoid apes. Cortex and Ganglia. Little is known of the particular order and manner of development in either the cortex or ganglia of the
cile
cerebellum.
As
to the cortex,
it
multiplication and development of the cells in the mantle layer, some of whose processes descend to the cerebellar ganglia and
Deiters's nucleus, and, (2) of the formation of contacts with in-
from pons, medulla and spinal cord. Corpus Restiforme of Cerebellum. Fibers from the cerebellar cortex descending to the medulla, and ascending fibers from the
growing
fibers
cord and medulla in their course to the cerebellar cortex, give rise to the corpus restiforme in the third month. Fibers from
the nucleus pontis
produced by
form the brachium pontis the brachium conjunctivum fibers from the nucleus dentatus.
fastigii
fifth
and nucleus
month
and 122). The pons develops simwith the cerebellum. The ventral zone of the metultaneously The neuroblasts formed therein encephalon thickens greatly.
(Figs. 118, II, I, 121
Pons
?)
and
fifth,
sixth
and seventh
pairs of cere-
lamina
cephalon, produces the neuroblasts which form the superior olivary nucleus and the superior part of the terminal nucleus of the trigeminal and of the acustic nerve. From the nucleus pontis axones
bellum.
ascend through the lateral walls of the metencephalon to the cereThey form most of the brachia pontis. At the same
time, about the fourth
tracts
composing the
tracts
basis pedunculi
grow downward
and intermediate
422
The pyramidal fibers to motor nuclei of down through the pons, intersecting its trans-
MYELENCEPHALON.
The myelencephalon forms
I,
the
medulla oblongata
and
122).
It is constricted off
twenty- eighth day; but, later, that constriction largely disappears and the common cavity of the two vesicles, broad in the middle
V
Fig. 125.
mm.
(McMurvz.
dz.
Dorsal zone.
fs.
Tractus
solitarius.
I.
Lip.
rp.
Roof -plate,
Ven-
and contracted
the
fifth
mature brain.
is
brain vesicle
by a deep furrow on the ventricular surface of each lateral wall (Figs. 125 and 126). That lateral furrow actually persists in the sulcus limitans and inferior fovea of the fourth ventricle.
dorsal zone
and a
myelencephalon stretches out widely and remains a single layer of epithelial cells. It forms no nerve tissue. Other portions of the myelencephalon develop
The
But by the expansion of the roof, just mentioned, quite uniformly. the dorsal extremities of the lateral walls are pushed outward and
MYELENCEPHALON.
forward almost to the plane of the
floor,
423
and a transverse section form of a very broad with the roof epithelium stretching between the two
letter into
a triangle.
Transverse sec-
tion through the lower half of the myelencephalon at the third or fourth week shows an elongated ellipse with a dorso-ventral
major
axis.
Internal Surface (Fig. 125). The lateral wall, in both upper and lower regions, presents the longitudinal groove which sepaThe median ventral groove rates the ventral and dorsal zones.
persists throughout
and the
by the
jar
Fig. 126.
Transverse section of the medulla from an embryo of eight weeks. (McMurrich after His.)
sf.
of medulla,
the trigeminal nerve, jr. Substantia reticularis. 01. Olivary nucleus Tractus solitarius. tr. Restiform body. XII. Hypoglossal nerve.
sulcus limitans and inferior fovea of the fourth ventricle (Fig. 96). situated between the median and each lateral
groove becomes the eminentia medialis. External Surface (Figs. 125 and 126).
On
face of the myelencephalon, along the ventral border of the dorsal zone, there appears very early an oval bundle of de-
scending fibers, called the solitary tract. It is composed of axones from the geniculate, glossopharyngeal and vagus ganglia, and constitutes the root-fibers of the intermediate, the ninth
and tenth pairs of nerves. At about the same time axones from the semilunar ganglion form a bundle of descending
424
fibers
They
Soon after the constitute the spinal tract of the trigeminal nerve. appearance of the solitary tract and the spinal tract of the fifth
nerve, the posterior margin of the dorsal lamina is folded outward and forward until it rests upon the external surface. That
fold,
which
is
above bundles
and places them in the position they occupy in the mature brain. Between the rhombic lips the roof epithelium stretches across the ventricle and pushes a transverse fold into it. The mesoblast which dips into that fold develops the chorioid plexus
of fibers
of the fourth ventricle (Fig. 91).
TABLE
X.
DERIVATIVES OF MYELENCEPHALON.
(Modified from McMurrich.)
Roof-plate
Roof epithelium
Nuclei funiculi
gracilis,
and
cuneati,
n.
tri-
and nucleus
tractus
spinalis
Dorsal Zone
-]
Arcuate nuclei
and tenth)
Sensory tracts grow up through
it.
Nucleus
lateralis inferior
and eleventh
(its
bulbar
and
tract
twelfth
Motor
through
it.
Surrounding Mesoblast
Meninges
Chorioid plexus of fourth ventricle.
The substance
layers
at
of the myelencephalon, like other divisions under the microscope three distinct
the fourth
week
of embryonic
life
(Fig.
125).
The
MYELENCEPHALON.
425
outer layer, the marginal velum, is composed of neuroglia; the middle, or mantle layer, of neuroblasts; and the inner, or ependy-
mal
layer
is
made up
of
columnar
epithelial cells.
ciliated
The
The
cells of
become
lining
of the ventricle.
mantle,
middle
layer
Its neuroblasts
nerve nuclei and other nuclei of the medulla oblongata. In the dorsal zone the neuroblasts form the terminal nuclei for the eighth,
ninth and tenth cerebral nerves and the nucleus funiculi
gracilis,
nucleus funiculi cuneati and nucleus of the spinal tract of the trigeminal nerve. As early as the fourth week axones may be traced
gracilis
and nucleus
funiculi cuneati,
ventro-medially, toward the point where they very soon form the Neuroblasts which have wandered from the dorfillet decussation.
sal
zone form the arcuate, the olivary and accessory olivary nuclei.
olivary nuclei are developed quite late in the intrauterine
(sixth
The
life
month).
The
nucleus of the spinal tract of the n. trigeminus, the olivary and arcuate nuclei are all products of the rhombic lip (Cunningham). From the neuroblasts of the ventral zone are developed the gray matter and fibers of the substantia reticularis alba et grisea, and
the motor nuclei of the twelfth, eleventh (cerebral part), tenth and ninth pairs of cerebral nerves.
The neuroglia layer, or marginal velum, forms the supporting matrix for the tracts of fibers in the medulla. By the third month the funiculus gracilis and funiculus cuneatus, extensions
of the
same
The
restiform
body
is at
developed, and the tracts of the lateral area of the medulla are visible. The medial longitudinal bundles appear near the median
raphe in the ventral zone at about the same time; and, ventral to them, fibers from the fillet decussation insinuate themselves and
form the interolivary stratum of the medial fillets. The great motor tracts from the anterior central gyrus of the cortex reach the medulla at the fourth month. Growing downward in the
neuroglia layer, on either side of the median
line,
426
medial
of the
medulla oblongata.
The form
of the
SPINAL CORD.
That portion
of the neural tube
is
which
is
the embryonic spinal cord (Figs. 16 and 118). metencephalon It is of nearly uniform size from cephalic to caudal end. The lumen of the neural tube in this region is at first large and elliptical in
shape. Later, at the sixth week, it has a diamond shape, the acute angles of the diamond being formed by the roof-plate and floor-plate of the canal; it is lined with columnar ciliated
cells
(Fig.
117).
As
it
is
contracted
until
The
to
canal
is
form the ventriculus terminalis in the filum terminale internum The spinal part of the neural tube forms the whole (Fig. no).
substance of the spinal cord, with the exception of the great motor
tracts that
grow
into
it
of embryonic life
it from the brain, and the sensory tracts and from the spinal ganglia. At the sixth week the anterior and posterior roots of the spinal
nerves are clearly seen; they are horizontal in direction. cord extends to the fourth mesoblastic somite of the coccyx,
the somites are
The when
first laid down; but, as no neuroblasts are developed the three lower segments of the human cord, they form a conby nective tissue strand, the primitive filum terminale (McMurrich). The cord occupies the entire length of the spinal canal until the third
month, when the caudal end begins to recede and the filum terminale to lengthen. It reaches only to the third lumbar vertebra at birth and, in the adult, but to the lower border of the
first
lumbar vertebra.
With the rapid growth of the spinal column, the roots of the lumbar, sacral and coccygeal nerves and the filum terminale become greatly elongated and form the cauda equina.
The investing mesoblast of the neural tube develthe meninges of the spinal cord. ops Zones (Figs. 117 and I27A). By the sixth week of intrauterine
Meninges.
SPINAL CORD.
life
427
and a
the neural tube, in the spinal region, is divided into a ventral dorsal zone by a lateral groove on either side, continuous
The
with that formed in the rhombencephalon two weeks earlier. tube presents externally, opposite to each lateral groove, a furrow in the mantle layer representing the concavity of the gray crescent and called the central
fissure;
it
is
occupied later
by the lateral pyramidal and other tracts. The whole of the spinal cord ventral to the bases of the posterior columnar of gray matter is represented by the ventral zones. From them grow out the
<fA
Fig. 127.
Transverse sections through the spinal cords of embryos of (A) about four and a half weeks and (B) about three months. (McMurrich after His.)
dh. Posterior cB. Fasciculus cuneatus (Burdachi). cG. Fasciculus gracilis (Golli). columna. dz. Dorsal zone. fp. Floor-plate, ob. Oval bundle (Hisi). rp. Roof -plate. vh. Anterior columna. vz. Ventral zone.
In the dorsal zones the poscolumnae and the posterior columns of fibers are developed.
receive the posterior roots of the spinal nerves (Figs. 117,
They
somewhat
Three Histologic Layers (Figs. 116 and I27A). At a time earlier than the division into ventral and dorsal zones,
fifth
428
presents three microscopic layers, like those seen earlier in the brain vesicles.
outer layer, or marginal velum, composed of neuroglia, thin except in the central fissure and in the floor and roof very In those situations of the tube on either side of the median line.
is
The
funiculi of
the cord.
The
blasts.
mantle
layer
is
gray matter.
sixth
At the fourth or
all
week
it
nearly
Pbsterwr
Fig. 128.
Mode
of origin of anterior and posterior roots of spinal nerves. matic. (Brubaker and Edinger after His.)
Diagram-
stance
is
blasts develop
They
median
line,
the an-
columna.
The axones
of the
more
centrifugally out of the antero-lateral surface of the neural tube. They form the anterior roots of the spinal nerves.
blasts
grow
The more
on account
dorsally located neuroblasts develop axones also; but of meeting resistance, according to His, they become
longitudinal and form the fasciculi proprii of the cord and, perhaps, the ascending anterior cerebello-spinal and the spino-thala-
SPINAL CORD.
429
mic
It
tracts.
in the dorsal
zone develops
later.
becomes transformed into the posterior columna; its neuroblasts form largely intrinsic neurones, their processes remaining in the
spinal cord,
tract
others
give
rise
to
the
posterior
cerebello-spinal
which ascends through the marginal velum to the vermis superior of the cerebellum. Axones from the spinal ganglia,
forming the posterior roots of the spinal nerves, grow into the dorsal zone; and each axone divides T-like into a descending and an
ascending branch, and also gives
off.
many
collaterals
both before
and
after
division.
The ascending
fibers for
;
oval bundle of His on the surface of the tube later, they constitute the marginal tract (Lissaueri) and the fasciculus cuneatus (Burdachi) and fasciculus gracilis (Golli).
The third, or ependymal layer, lines the ventricle. It is composed of neuroglia covered internally by columnar ciliated cells, which appear about the fifth week.
The Longitudinal Tracts
(Figs. 101, 102
and
103).
The
white-
columns of the spinal cord are formed in the superficial, or neuAt first they are composed of nonmedullated fibers, roglia layer.
and
it
is
myelin sheaths
at definite periods between the fifth and ninth months of intrauterine life (Flechsig). The anterior nerveroots and the anterior and lateral fasciculi proprii, and the posterior nerve-roots
of
His are
first
to
make
tracts
their appearance.
The axones
of these nerves-roots
and
may be
The
ascending cerebello-spinal
tracts follow,
fourth or
fifth
month.
According
to Kahler, as
quoted by Cun:
ningham, the tracts of the spinal cord are medullated as follows 1. Fibers of the fasciculus cuneatus with the posterior nerveroots,
(fifth
2.
month).
of
Fibers
anterior
and
(fifth
and
sixth months).
3. 4.
Fibers of fasciculus gracilis (sixth month). Posterior cerebello-spinal tract (seventh month).
430
5.
tracts
month).
fissure
fissure.
is
Pyramidal tracts (ninth month). Fissures (Fig. 101). The so-called posterior median At no time is it a true in reality a septum of neuroglia.
6.
Its
mode
of formation
is
still
in doubt.
It
appears to be proof
to the elongation
ependymal
cells.
From
it
the sixth
week
this neuroglia
septum
ex-
The posterior lateral sulcus is the groove between the lateral border of the posterior column and the dorsal border of the It is the development of the posterior and lateral lateral column.
columns that produces the fissure. The embryonic central fissure is obliterated and the postero-lateral surface of the cord rendered prominent by the formation of the lateral fasciculus proprius,
the cerebello-spinal and the pyramidal tracts. Anterior Median Fissure. That is a true fissure.
to
It
begins
be formed
when
It
the earliest fibers of the anterior fasciculus proprius are developing. deepens with the growth of the gray columnae and of the
anterior fasciculus proprius
and
its
amidal
tracts.
In
anterior surface
by the descent of the anterior pyrmanner there is produced a bulging of the on either side of the median line, which increases
this
with the medullation of the longitudinal tracts up to the ninth month. The ridges thus produced, failing to fuse completely,
become the
median
fissure.
The
partial
fusion which does occur between the two ridges is due to the formation of the white anterior commissure of the cord.
ADDENDA.
According
distinct
tion,
to Head, Rivers and Sherren (Brain, 1905) three mechanisms operate in the production of common sensaone concerned with deep sensibility and two with superficial
sensibility.
The mechanism
of
supplying muscles, tendons, joints and ligaments. Though it may be affected by environment, it is chiefly acted upon by stimuli
originating within the organism, such as pressure (resulting from weight and inertia) and tension. It may give rise to painful
sensations, but its principal concern is with muscular tonicity and equilibrium. The nerves of the deep mechanism constitute " " the of Sherrington. They are so named beproprio-ceptors
itself
The mechanisms
by Head, Rivers and Sherren the Protopathic The protopathic mechanism responds to painful cutaneous stimuli and to extremes of heat and cold. It may set up reflexes without arousing consciousness. Cutaneous localization is no part of its function. The mechanism of epicritic
sibility are called
and the
Epicritic.
sensibility is
to coolness.
By
its
adapted to mild stimuli, to light touch, to warmth, it the localization of cutaneous stimuli is accomactivity usually involves consciousness.
It
plished
and
may
and speaks of these two mechanisms as the extero-ceptors 'gives the name "intero-ceptors" to the nerves of the interior surfaces, those of the alimentary tract, etc.
"
Sherrington "
The
"intero-ceptors"
and give rise to digestive reflexes; but they may also produce pain and temperature and may respond to stimuli occasioned by tension and sensations,
are especially adapted
to chemical stimuli
pressure.
INDEX.
Abducent nerve,
45,
153, 321
Accessor}' lemniscus, 227 nerve, 5, 49, 153, 293, 322 nucleus funiculi cuneati, 311
338
lateral sulcus of medulla, 41,
153,
295
longitudinal bundle, 158, 161, 163, 219, 276, 295, 297, 305,
226,
363 sympathetic fibers, 348 After-brain, 33 Ala cinerea, 149, 270, 290, 317 Alveus. 200, 20 1
root,
34i, 345, 355 median fissure of cord, 335, 337. 349, 430 median fissure of medulla, 41, 285, 286, 295, 337 median vein of medulla, 24
orbital gyrus, 74 parolfactory sulcus. 86
,
341,
63, 67
345, 353
radicular artery,
foot-line,
Ansa
lenticularis, 100,
145
331
peduncularis, 145, 212 Anterior area of medulla, 293 association center, 182 brain vesicle, 30 calcarine fissure, 89 central gyrus, 39, 59, 60,
335, 338
root of spinal nerve, 327, 334, 349, 3 6 3. 399, 427. 428 slender lobule, 252, 253, 256
spinal artery, 5, 15, 24, 27, 330,
61
33i
subarachnoid space,
central artery, 331 cerebral artery, 4, 6, 15, 16, vein, 23 chofioidal artery, 15, 1 8, 19
25
109,131,136,137,234,235,
columna, 335, 339, 399, 427,
243
Antero-lateral ganglionic arteries, 15,
l8,
19, 22
428
neurones, 165, 340 external arcuate fibers, 292, 295,
297
fasciculus proprius, 354 horn of lateral ventricle, 120,
fasciculus proprius,
290
137,
209, 235
inferior cerebellar artery, 15, 27,
29
internal frontal arteries, 16
quarti (Key and Retzii), 9, 290 Aqueduct of cerebrum, 37 Aqueductus cerebri, 37,75,152, 161,
lateralis
ventriculi
270
433
28
434
Arachnoid granulations,
of brain, 8, 9 of cord, 326, 327,
5,
INDEX.
6,
65
tract,
361
329
265
326
fibers of
Arbor
vitae,
247, 262
Auditory artery, 5
center, 70, 71
conduction path, 160 paths, 280, 385 Auditory and facial nerve, 5
Area
149, 270, 290, 318 (Brocae), 47, 75, 91, 94, 200 postrema, 270, 318
parolfactoria
for educated
acustica,
Aula
of
third
ventricle,
30,
137,
141
Axones, 168, 169, 398, 400
Axone
hillock, 167,
169
movements
of cere-
brum, 178
Arteria basilaris, 14
carotis interna, 14, 25 cerebelli inferior anterior, 27, 29 inferior posterior, 27, 29
25
75
media, 16
posterior, 17,
25 25
chorioidea anterior, 18
communicans
anterior, 14,
posterior, 14,
25
6,
US,
7
27, 330
meningea media,
259
Bechterew's accessory lemniscus, 227
superior nucleus, 277, 280, 308 Betz, giant pyramids of, 188 Bipolar neurones, 400, 401 Biventral lobules, 252, 254, 255 Blood supply of brain, 14
of cerebellum, 27 of cerebrum, 14 of medulla oblongata, 24 of pons varolii, 24 of spinal cord, 330 Body of fornix, 101, 109 Boundaries of fourth ventricle, 313, Brachia of corpora quadrigemina, 30,
146, 164
331
Ascending
anterior
287,
cerebello-spinal
tract,
anterior
22
Brachium conjunctivum
brachia
cerebelli (or
13,
246,
252, 261,
275, 289
INDEX.
Brachium
inferius of orpora quadri-
435
gemina,
146,
pontis,
246, 247, 252, 255, 264, 267, 289 superius of corpora quadrigemina, 147, 157, 163, 164, 207, 416, 418 Brain, 31
general considerations, 34 vesicles, 30, 34, 395, 402, 403
Centermedian (Luysi)of thalamus,2n of abstract concept, 182, 182 of concrete concept, 182, 183 of crescent, 344
of equilibrium, 71, 181 of intonation, 71, 181 of macular vision, 67 of muscular sense, 178 of optic reflexes, 324 of orientation, 71, 181 of smell, 82, 94, 181 of stereognosis, 178
Bruce
and
Muir's
361
septo-marginal
lateral
tract,
of taste,. 94, 98, 181 of visual memories, 67, 69 Central canal of cord, 399, 428 of lower medulla, 305 cells of anterior columna, 343,
345
CajaPs moss-like appendages,
260,
259
tassel cells,
199
gray substance of medulla, lobule, 247, 250 part of lateral ventricle, 112
305
Calamus
133,
410
Calcarine artery, 17
fissure,
Calleja's olfactory islets, 199 Callosal sulcus, 80, 86, 91, 95,
410
Calloso-marginal sulcus, 86 Canalis centralis spinalis, 337
Centrosome, 167
Capsula interna, 99
Cauda
equina, 333,
334
Caudate nucleus, 101, 105, 109, 115, 116, 123, 125, 137, 145, 204, 207, 209, 235
265,
292,
295,
3"
(poste<
-
Cava subarachnoidealia, 8
Cavernous
sinus, 3,
Cavum
septi pellucidi,
345, 347
Cells of anterior columna, 340 of granular laver of cerebellum,
Cerebellum, 31, 33, 35, 36, 37, 3 8 43, 53, 209, 242, 418, 419 blood supply of, 27, 29, 30 veins of 29, 30 Cerebral aqueduct (Sylvii), 4, 30, 34
258
of posterior columna, 347 of Purkinje, 257, 258, 259 Cell and fiber lamination of cerebral
cortex, 183,
87, 132, 147, 152, 247, 417 and medulla, 177, 411 gray substance, 174 hemispheres, 30, 35, 37, 51, 98
cortex
localization, 177
peduncle, 209,
reflexes,
267
197
392
43 6
Cerebro-pontal paths,
frontal,
INDEX.
temColliculi
reflexes.
Cerebrum,
30,
of corpora quadrigemina, 147, 149, 163, 243, 246 Colliculus facialis, 149, 269, 270, 277, 3i7 inferior of corpora quadrigemina, 149, 16,3, 220
333flexures,
334, 335
402
Chiasma opticum,
Chief
nucleus
of
38, 82,
83
nerve
vestibular
149, 159, 163, 219, 243, 416 Columna anterior, 339 fornicis, 105, ITO, 131, 239 lateralis, 335, 341, 344
ina,
(Schwalbe), 283
Chorioid epithelial lamina of lateral ventricle, 124, 412 plexus of fourth ventricle, 13, 290, 314, 424
of lateral ventricle, 10, 80, 90, 120, 124, 125, 127, 136,
Columnaeof
Comma
no, 131, 132, 137, 139, 209, 212, 235 tract of Schultze, 341, 361 Commissura anterior alba, 351*!
anterior cerebri, 31, 136, 201 anteria grisea, 350 grisea of spinal cord, 338, 350
209, 417
tela of fourth ventricle,
habenularum, in, 135 hippocampi (Fornix), 51, no, 125, 131, 236, 412, 414
inferior (Guddeni), 218, 221
10
n,
90,
95, 131, 132, 135 Chorioidal fissure, 80, 86, 90, 95, 101,
124, 127,
posterior cerebri, 132 posterior of cord, 350 superior (Meynerti), 218, 221 Commissural fibers of cerebellum,
262, 265 of cerebrum, 224, 234 nucleus, 305, 306 Common sensor}' area of cerebrum, 178
147
vein,
n,
21,
129
204
Chromophilous
cells,
paths 375, ^77, 379 Confluens sinuum, 2 Con us medullaris, 31, 334
390
Convex surface of fore-brain, 52 Corona radiata, 209 Cornu anterius ventriculi lateralis, 120 commissural tract, 345, 362
inferius ventriculi lateralis, 123,
133
Claustrum, 101, 145, 204, 209, Clava, 13, 246, 289, 300, 310 Cochlear nerve, 280
nucleus, path, 386 Cohnheim's end
235
133
Corpora albicantia
(see corpus milla), 85, 221
mam-
18
Collaterals, 168, 169, 398 Collateral fissure, 75, 80, 81, 86, 90,
geniculata, 30 quadrigemina, 28, 30, 37, 109, 146, 163, 219, 418 mamillaria (see corpus mammillare)
INDEX.
Corpus callosum, 6, 25, 30, 51, 87, 91, 101, 104, 109, 117, 112, 145, 209, 234, 235, 411, 413. 414
dentatum, 260
geniculatum mediale, 143, 147, 149, 218, 221 geniculatum laterale, 143, 147, 218, 416
Luysi, 143,
Crista
galli,
437
4
Crossed
145
(or
mammilare
corpora
mam-
47, 75, 82, 85- 87, 95, 131, 145, 153, 2O7, 2O9, 221, 267, 417 medullare cerebelli, 242, 244, 247, 255, 261, 262 pineale, 30, 75, 132, 137, 243 restiforme, 149, 246, 264, 288,
millaria), 30, 38,
pyramidal tract, 287 of hippocampal gyrus, 196 Cms fornicis, 95, 101, 105, no, 124, 131, 133, 237 Oilmen monticuli, 87, 243, 245, 247, 250 Cuneate gyrus (see cuneus) tubercle, 13, 289, 310 Cuneus, 25, 97
Crown
Declive monticuli, 87, 243, 247, 251 Decussatio lemniscorum, 291, 292
301, 421 striatum, 28, 36, 37, 98, 115, 204, 410
275
Deep
of pyramids, 209, 267, 291, 294 transverse fibers of pars basilaris pontis, 271, 275 veins of pons, 27 vena cerebri media, 23
259
of cerebellum, 421 Cortical areas of convex surface of
and ganglia
Defecation
179, 183
of smell, 82, 94, 181 gray matter of cerebrum, 177 connection of abducent nucleus,
of Nissl, 171 of Waller, 171 Deiter's cells, 340, 348 dorso-lateral vestibular nucleus,
282
of anterior columna, 343 of cochlear nuclei, 310 of facial nucleus, 282 of genetic nuclei of cerebral nerves, 324 of motor nuclei in medulla, 305 of nucleus alae cinereae, 306 of oculomotor
nuclei, 223
280, 308 Dendraxone, 168 Dendraxones of Golgi, 191 Dendrites, 165, 167, 169, 399, 401 Dentate fascia, 82, 101, 124, 127, 415 gyrus, 411, 414 nucleus, 255, 260, 277
Derivatives and ventricles of brain
vesicles,
and trochlear
403
of dienccphalon, 415 of mesencephalon, 417 of metencephalon, 419 of myelencephalon, 424 of telencephalon, 404 Descending anterior cerebello-spinal tract, 287, 298, 305, 311, 34i, 345, 35 2 356 part of medial longitudinal bun-
nerve, 324 of trigeminal nucleus, 281, 307 of vestibular nuclei, 308, 323 fillet, 104, 160, 216, 232, 233
localization, 177 system of arteries, 15 Corticifugal fibers of cerebrum, 225
438
INDEX.
Embryological
tracts,
method
352
of
locating
cord,
352
Destruction
389
Destructive lesions of cerebral cortex,
172
Eminentia cinerea, 317
collateralis, 93,
183
124,
131, 133,
411
medialis, 290, 423 Emissive motor area of cerebrum, 63. 177 Encephalon, 31 general considerations, 34 End arteries of Cohnheim, 18 End-brain, 30, 33, 50, 51, 73 End-brush of axones, 168
335
3i. 357 motor paths, 367 pyramidal tract (see anterior pyramidal tract)
Entry zone, 341, 345, 359 Ependymal cells, 173, 174, 397 layer of embryo, 398, 425, 429
Epiblast, 395, 396 Epiblastic sustentacular tissue, 173 Epiphysis, 135
Dorsal
accessory
olivary
nucleus,
297, 312
longitudinal fibers of pons, 272 tegmental decussation (Meynerti), 156, 157, 159 view of brain ventricles, 141
of
locating
zone,
422, 427
cells of cord, 340,
Dorso-medial
341
280, 307
Dorso-lateral
cell-group,
101, 105, 113, 145, 209, 235 layer of large pyramids of cerebrum, 184, 187
capsule,
341, 343,
345
vestibular nucleus (Deitersi),
280,
308
Dorso-ventral fibers of medulla, 292 of spinal cord, 351
Dura mater
Facial nerve, 45, 153, 209, 321 Facies anterior of medulla, 286
cerebelli inferior, 251
329
7
326
of brain
medulla, 287
95
Falx
cerebelli, 2,
3, 4, 6, 407 Fascia dentata, 90, 97, 131, 200, 201 Fasciculus antero-laleralis proprius,
cerebri,
i,
2,
fibers, 344 Elements of olfactory bulb, 196, 203 Embryologic divisions of the brain table, 30
3H, 341/345,
INDEX.
Fasciculus
cerebello-spinalis
439
poste-
rior, 301,
357
411
75, 80, 81, 86, 90, 131 hippocampi, 80, 86, 90, 95, 131,
collateralis,
358 cuneatus, 335, 341, 361, 427 descendens cerebello-spinalis anterior, 298,
lateralis,
2OI
longitudinalis
cerebri
35,
39,
356
427
43, mediana
52, 53 anterior,
337 409
posterior,
337
longitudinalis in ferior,
131, 201,
354
237, 240
inedialis, 156, 276, 296,
occipito-parietalis, rhinalis, 93
cerebri, 35, 55
transversa cerebelli, 36
Fissural arteries, 331 Fissure, 52 Fissures of convex surface of cereof medial
pyramidalis, 148
superior, ventralis
237, 240
(see anterior longit-
341,345,
239,
brum, 52 and
85
tentorial surface,
359
occipito-fron tails
(Foreli),
241
olivaris, 208,
of spinal cord, 335, 337, 430 Flechsig's oval tract, 361 sensory bundles of internal cap
sule,
232
perpendicularis,
237, 241
(Vicq
d'
thalamo-mammillaris
Azyri), 85,
422,
Fastiguum, 244, 247 Felt-work of Kaes, 184, 191 Fiber zones, 184
Fibers of granular layer of cerebellar cortex, 260 of Meynert, 192 of superficial layer of
427
Folium vermis, 87, 243, 247, 251
cerebral
cortex, 257
156,
157, 159
159
decussation, 292,
Fore-brain, 30, 31, 35, 50, 51, 407 Formatio reticularis of cord, 335 of mid-brain, 155, 157, 159,
334
Fimbria hippocampi, 124
First cerebral vesicle,
161, 273, 339 of pons, 273, 275, 277 Fornix, II, 30, 87, 91,95,97, 101,
109,
35
61,
237,
Fossa cerebri
lateralis,
413
159
Qi
rhomboidea, 314
440
Fourth nerve,
ventricle,
3,
INDEX.
5, 13,
289
289, 313,
9,
242,
275,
422
45,
Fovea
Foveolae granulares, 6
Fnenuknn
Frenulum
of Giacomin,
94
Geniculate bodies (metathalamus),5T, 204, 207, 417 Gennari's or Baillargic line, 188
Genu
75, 87, 91, 98 part of internal capsule, 103 pole of cerebrum, 50, 53 stalk of thalamus, 103, 216, 226, 232 Fronto-marginal sulcus, 60 Fronto-pontal tract, 103, 151, 159,
73-.
.
capsulse internae. 103, 113 inferius of central sulcus, 56 internum of facial nerve, 283
Germinal
225,
226
109, 1 20, 121 area, 35 cells of neural crest, 172 of His in neural tube, 172
175, 182, 183 335, 352 cuneatus, 288, 300, 305, 311 gracilis, 288, 300, 305, 311
Globus pallidus of lentiform nucleus, 116, 145, 209, 235 Glomus chorioideum, n, 120, 125,
129
Glossopharyngeal nerve, 5, 46, 153, 209, 293, 320, 321 Golgi cells, 165 of spinal cord, 165, 340, 347 dendraxones, 191 Goll's column, 300, 360
Gordinier's writing center, 175, 178, 182
lateralis, 335,'
352
335, 35 2
separans,
layer, 191
270
79, 81,
Fusiform gyrus,
95
Gowers's tract, 276, 356 Granulationes arachnoideales, 5 Granular layer of cerebellar cortex
258
Gambault
and
Philippe's
129 median
Granule
Gray
cerebellum, 258, 259 anterior commissure, 335, 350 commissure of cord, 335, 338,
cells of
35
crescent of spinal cord, 339 matter of cerebellum, 256
of cerebrum, 164 of medulla, 302
151
267
spinale, 364 Ganglionar gray matter of cerebellum, 260 of cerebrum, 204 (Bevin Lewis), Ganglionic cells
of pons, 278 of spinal cord, 338 of stratum nucleare, 278, 302 radiatio Gratiolet's occipito-thala-
191
341
Genetic nucleus,
i.,
42, 319,
363
6,
INDEX.
Gudden's
inferior
441
vesicle, 405, 409 of cerebellum, 41, 242, 245,
commissure, 218,
Hemisphere
221,
235
252,
Gustatory center, 179, 181, 183 paths, 389 radiations, 232 Gyri breves insuloe, 65, 72 insular, 65, 72, 235 occipitales laterales, 68 superiores. 68
temporales transversi, 70 transitivi, 64 of convex surface of cerebrum,
59 of medial and tentorial surface, 93 Gyrus angularis, 61, 63, 67
centralis anterior, 59, 60,
419 of cerebrum, 35, 39, 43, 47, 53, 405 Hemispheria cerebelli, 242, 245, 252 Hiatus (Sylvii), 99, 115 Hind-brain, 31, 33
Hippocampal
fissure, 80, 86, 90, 91, 95, 101, 201, 409, 411,
416
gyms,
region,
81, 94,
95
201
80, 90, 101, 113, 131, 133, 145, 409,
Hippocampus,
124,
{
61
.
414,
416
posterior,
cinguli, 93, cortex, of
His
(see cal car avis), 125, 133 (germinal cell in neural tube), 172
minor
Histologic layers of neural tube, 427 of spinal cord, 397 Hoche's bandalette, 362, 345 Horizontal anterior ramus of lateral
fissure 57,
61
medius, 59. 61
superior, 59, 61, 95, 97 fusiformis, 75, 81, 97 hippocampi, 79, 81, 93, 94,
95,
97
lingualis. 8t,
Hypoblast, 396 Hypoglossal nerve, 5, 49, 153, 293, 322 nucleus, 295, 303
triangle, 317 Hypophyseal region, 37 Hypophysis cerebri, 37,
75
38,
47,
82.
74
84,
267
medialis 74,
75
64 75, 95, 97 semilunaris, 82, 94 subcallosus, 75, 87, 91, 97, 107, 200, 414 supracallosus, 107, 414 supramarginalis, 61, 63, 67 temporalis inferior, 61, 70, 71,
Indifferent
cells,
397
Indirect
75
medius, 61, 70, 71
superior, 61, 70, 71
72
235
efferent veins of pons, 27 external cerebellar veins, 29 frontal artery, 19, 22
298,
305,
fovea,
270,
290, 317
442
INDEX.
Internal layer large pyramids of cerebral cortex, 184, 1 88
57, 59 gray commissure of fore-brain, 84 horn of lateral ventricle, 101, 105, 123, 133, 141, 145
internal frontal artery, 19 lamina of internal capsule,
99,
sulcus, 39,
332
veins of cerebrum, 21
Interparietal sulcus, 39,
43, 57,
63,
414
Interpeduncular fossa, 145, 146, 153,
parietal lobule, 61, 63, 64 peduncles of cerebellum, 246, 264 of thalamus, 100, 145
36, 37, 95, 112, 128, 235, 405 Inhimescentia cervicalis, 333, 334,
335
lumbalis, 334, 335 Inverted pyramids of Martinotti, 191 Island (of Reil), 71, 74, 75,98, 105,
precentral
sulcus,
414
Isthmus
gyri fornicati, 94 rhombencephali, 28, 33, 418,
414
13, quadrigeminal colliculus, 147, 149, 163, 204, 207, 220, 418
sagittal sinus, 2, 3 semilunar lobule, 252, 255,
419
Jugular ganglion, 49
Kaes, medullation late in life, 412 Retzii, apertura lateralis ventriculi quarti, 9, 290
affixa,
255,
420
surface of cerebellum, 251,
of island, 74 striate veins, 23
Key and
252
Lamina
119
124
cinerea terminalis, 30, 36, 38, 82,
83.
Infundibulum,
-
109,
131,
109.'
J3 1
290
Interlobar boundaries, 55 Intel-mediate nerve, 45, 153, 320 olfactory stria, 77, 78
159, 225, 226 column of cellbodies, 341, 344 Internal capsule. 99, 101, 105, 109, H3, US, 145, 209,226,
tract ico, 148,
341,
Intermedio-lateral
345
column of medulla, 287 columna of cord, 335, 344
fasciculus
proprius,
287,
298,
412,-
417
354
19
fillet,
274-
INDEX.
Lateral fissure of cerebrum (Sylvii),
443
53,55- 57,61,63,91,414 geniculate body, 140, 143, 157, 204, 207, 218
longitudinal stria (see longitudinal stria) nucleus of thalamus, 211
orbital arteries, 16
gyrus, 74
cerebrum, 93
of superior surface of cerebellum,
373 pyramidal
341,
345, 358
recess of fourth ventricle, 289 sulcus of mid-brain, 159, 243
in,
Layer of fusiform
tex,
184 of large pyramids, external, 184, 187 of large pyramids, internal, 184, 188
of of
250 Lobules of cerebellar vermis, 247, 249' 253 Lobulus biventer, 245, 254 cen tralis, 87, 245, 250 gracilis, 245, 256 paracentralis, 64, 97 parietalis inferior, 63, 64 superior, 63, 64 Lobus centralis cerebelli, 250 culminis cerebelli, 243, 250 declivis cerebelli, 243, 250 folii vermis, 243, 251 frontalis. 59
limbus, 8 1 linguhe cerebelli, 250
noduli, 253
medium
cortex, 184, 187 of stellate and polymorphous cells of cerebral, cortex, 184, 188
Law
of Waller, 353
lateralis, 160,
Lemniscus, 159
parietalis,
274
Locus
Long
Lesions of anterior columna, 344 of corpus striatum, 208 of gray substance in cord, 350
of special sense paths. 389
300
of spinal cord, 352
fiSsure of
in
52,
strife,
cerebrum, 35, 39, 43, 53, 209, 235 medial and lateral, 97, 107,
Limbic
87, 91,
79
93, 97,
98
Limen
444
INDEX.
of spinal cord,
9,
9, 10,
Luys, center median of thalamus, 211 nucleus hypothalamicus of, 143, 151, 218
Gambault
and
Phillipe), 361
Lymph
Medulla oblongata, 24, 31, 33, 35, 36, 37, 38, 87, 242, 284, 422 spinalis, 31, 333, 334, 335 Medullary body of cerebellum, 242, 244
groove, 396 lamina?, 262
plate,
Mammillary bodies, 38, 51, 75, 85 Mantle layer, 398, 425, 428
Marginal gyrus, 95, 97 sinus, 5
tract,
ridges,
striae,
395 395
Membraaes,
341,
Lissauer's,
345,
359
velum. 397, 397, 398, 425, 428 Margo-infero-lateralis, 52, 75
Margo
75
327
75
39, 52, 75 supero-medialis, 39, 52 Martinotti's inverted pyramids, 191 Ma ssa intermedia, 95, 137, 139, 145, 221, 222, 417
Mesencephalic flexure, 402 Mesencephalon, 28, 30, 31, 144, 402, 417 Mesoblast, 396 Mesoblastic sustentacular tissue, 174 Metathalamus, 140, 143, 218
Metencepbalic flexure, 402 Metencephalon, 28, 31, 266, 402, 418
33,
Medial
accessory
295.
297,
36,
cerebral veins, 22 column of cell-bodies, 340 fillet, 157, 159, 160, 161, 231,
273,
.
Methods
35 2 Meynert's association
236
295, 297, geniculate body, 140, 143, 146, 147, 149,' 157, 159, 204,
35
157, 159
solitary cells, 195
207, 218
longitudinal bundle, 156, 157, 159, 161, 275, 276, 277, 295, 296, 297, 305, 311,
75
Mid-brain, 30, 31, 33, 36, 37, 5o,;87, 144, 243, 417 Middle association center, 183 brain vesicle, 30 cerebral artery, 4, 15, 16 commissure, 137, 222 frontal gyrus, 39, 59, 61 sulcus, 57, 60 internal frontal arteries, 16
gyrus, 74, 75 ponto-spinal tract, Collieri, 158, 297. 355- 373 ; surface of fore-brain, 92, 95
meningeal artery,
3,
5,
6,
and
47,
75,
77-
78
70, 71
INDEX.
Midgracile sulcus, 421 Mixed ascending and tracts, 352
Neuroglia, 172, 399
.445
descending
cells,
173
Monro,
Neurolemma (Schwann),
Neurone 165,
167,
center, 167
168, 169
415 Moss-like appendages of Cajal, 259, 260 Motor area of cerebrum, 177 center for foot, 98, 178 cerebral nerves, 321
fibers of
cerebrum 225,
memory
34
neurone, 169
paths, 229, 366, 367, 371 roots of cerebral nerves, 321 root of spinal nerve, 363
169
297, 303, 304,
Nucleus
alae cinereie,
speech center, 178 Motorial end-plates, 168, 169 Myelencephalon, 28, 31, 33, 266, 284, 402, 422 Myelin sheath of axones and dendrites, 168
Myelinization, 412, 429
ambiguus, 297, 303, 304, 315 amygdala?, 113, 119, 199, 207,
239
arcuatus, 303 caudatus, 116
centralis inferior, 278, 279
Naming
corporis trapezoidei, 279 dentatus, 255, 260, 261, 277 dorsalis (Stilling! and Clarki),
348
dorso-lateralis (of Dieters), 283,
308
emboliformis, 260, 261, 277 262 fastigii, 261, 277, 260,
funiculi cuneati,
320
Nervus abducens,
45, 267, 321 accessorius, 49, 267, 322 acusticus, 46, 267, 320
310
gracilis,
321.
277
hypoglossus, 49, 267, 322 intermedius, 45, 267, 320 oculomotorius, 42, 267, 321
opticus, 25, 42, 83,
trochlearis, 45,
267, 320
intercalatns,
270 270
297, 303, 304
lateralis inferior,
267, 321
vagus, 46, 267, 321 Neural canal, 396 crest, 35, 172, 395, 400 groove, 35, 395 tube, 35, 172, 395 Neuraxones, 168 Neuroblasts, 398
209
282
446
Nucleus
vcstibularis,
INDEX.
270,
312 363
283, 305,
37- 315
olivaris inferior,
originis, 42, 319, 324,
Occipito-parietal fissure, 409, 411 sulcus, 17, 43, 53, 56, 57, 86,
89.
Qi, 95
radiation,
Occipito-thalamic
137,
213,
217,
226, 234
145, 153,
ruber, 217 terminalis, 42, 319, 365 tractus solitarii, 302, 305, 306
spinalis n. trigemini,
2Q5, 305,
159
3".
315
314, 3 6
281,
of anterior tubercle, 211 of abducent nerve, 277, of accessory nerve, 304, of Bechterew, 277, 308
of colliculus inferior,
203, 207
center, 97 cortex, 195 divert iculum,
315
161, 164,
220
of corpus mammillare, 85, 157 of descending root of trigeminal
nerve, 155 of descending root of vestibular nerve, 280, 283, 308 of external arcuate fibers, 302
of facial nerve, 277, 282, of fascia dentata, 201 of fourth nerve, 155, 315 of hypoglossal nerve, 303, of neurone, 167, 169 of oculomotor nerve, 152, of optic nerve. 315 of pulvinar, 208
315 315
315
36, 37 199 lobe, 73, 77, 98 nerves, 42, 320 path, 383 projection fibers, 232 sulcus, 74 striae, 75, 7 tract, 47, 75, 77, 78, 267 triangle, 75, 77, 79 Oliva, 287 Olivary fasciculus, 162, 208, 277, 298
islets (Calleja),
423, 425
pedicle, 279 Olive, 31, 41, 153, 287
Ophthalmic
280
of trigeminal nerve,
277,
281,
Optic center, 181 chiasma, 30, 38, 47, 75, 83, 95, 147, 153, 213, 235, 415 commissure (see optic chiasma),
I3 1 cup, 405 nerve, 4, 38, 42, 83, 153, 213,
315
of trochlear nerve, 152, 223, 315 Ninth, tenth and eleventh nerves, 3 Nissl bodies, 167, 169
degeneration, 171
320
path, 213, 384 radiation, 104,
232,
Nodes of Ranvier, 168, 169 Nodular ganglion, 49 Nodule of cerebellum, 245, 247, 252, 253 Nodulus cerebelli, 87, 253
Notochord, 396 Obex, 13, 270 Oblique fasciculus of pons, 267, 269 Occipital lobe, 53, 67, 75, 87, 91, 98
part of internal capsule, 103
pole, 50, 53, sinus, 2, 5
234
68
INDEX.
Origin of posterior root of spinal
447
Oval
tract (Flechsigi),
Pia mater of spinal cord, 329, 329 of brain and cord compared, 13 Pineal body, 36, 37, 51, 91, 95, 109,
131, 132, 137, 147, 157, 243, 416, 416 recess. 132, 416 stria, 135, 137, 239 (see stria
Pacchionian bodies, 5 Paracentral lobule, 64 Paramedial sulcus, 39, 57, 59, 60 Parietal lobe, 53, 63, 87, 91, 98 stalk of thalamus, 103, 104, 226
Parieto-temporal artery, 17, 22 Parolfactory area- (of Broca), 79, 86 Pars anterior rhinencephali, 77
basilaris pontis, 266 dorsalis pontis, 266, 273 frontalis capsulae internae,
medullaris thalami) Pituitary body, 84 (see hypophysis) Plexiform layer of cerebral cortex, 184
Plexus basilaris, 4
chorioideus
I2O,
ventriculi
lateralis,
2OI
103,
chorioideus ventriculi quarti, 290 venosi vertebrales interni, 332 Plica chorioidea of fourth ventricle,
H3
mammillaris hypothalami, 28,82, 221, 417
occipitalis capsulae internae, 103,
419
Pole of island, 71 Polus insulae, 71
Pons
(varolii),
24,
28, 31,
33,
37,
H3
optica hypothalami, 28, 82, 221,
38,87,101,145,153,242,
266, 418, 421 Pontal arteries, 24 Pontine flexure, 402 Ponto-spinal tracts, 280, 373
4iS
posterior rhinencephali, 77 Pathological method of locating tracts,
353
Path carrying
for tactile
visceral,
muscular and
381
sense,
tactile impulses,
420
of cerebrum, 63, 410 Postdeclivil sulcus, 243, 247, 249,
and muscular
375, 377
for tactile, pain
and temperature
420
Post-dorso-lateral
cells,
impulses, 276, 379, 381 through red nucleus, 370, 371 Peduncle of corpus callosum (see
Postnodular sulcus, 247, 252, 252. 420 Postparietal gyrus, 63, 67 Postpyramidal sulcus, 245, 247, 252, 253, 420 Posterior anastomotic vein, 23 area of medulla, 293, 300 association center, 182
brain vesicle, 33, 35
calcarine fissure, 89
central gyrus, 39, 61, 63, 64 cerebellar notch, 36, 243, 244
Permanent fissures and sulci of cerebrum, 409, 414 Perpendicular fasciculus, 237, 241
Petrosal
glossopharyngeal
ganglion,
46 Petro-squamosal sinus. 5
Physiological
tracts,
252
cerebello-spinal
fasciculus,
264,
method
353
of
locating
305,
3",
288,
341, 357
329
9,
of brain,
448
Posterior
INDEX.
commissure of cerebrum, 95, 109, 131, 132, 137, 416 of cord, 335, 350 communicating artery, 4, 14, 15 fasciculus proprius. 362 horn of lateral ventricle, 123,
Precuneus, 95, 97
Predeclivil
sulcus,
420
Preolivary nucleus, 278, 279
Prepyramidal sulcus, 245, 247, 252, 252, 420 Prevertebral ganglia, 401 Primary brain vesicles, 30, 35, 395 fissures of cerebrum, 409, 411,
413
Primitive streak, 35 Principle or chief vestibular nucleus
(Schwalbe's), 307 Processes of dura mater, i of neurone, 167, 169 Processus durae matris, i
338
internal frontal arteries, 16, lateral sulcus of cord, 335,
25
337
43
lateral sulcusof medulla, 41, 153, 286, 295, 297
149,
longitudinal bundle (see medial longitudinal bundle) median fissure of cord, 335, 337,
263
of cerebrum, 224
169
Psychic acustic center, 71, 181
center of abstract concept, 60,
perforated
substance,
38,
47,
182, 183
center of concrete concept, 182,
cms
183
area, 178 optic center, 181 sensory area, 178, 67 Pulvinarof thalamus, 140, 149, 211,
motor
207,
315.
428, 429
slender lobule, 252, 255
spinal artery, 24, 27,
394 394
331
Purkinje's
265
Putamen
Pyramid
of lentiform nucleus,
116
345, 399
of
column
tract,
(Burdach'i),
254
of medulla, 41, 153, 209, 267,
2 93. 2 94 295, 297, Pyramidal cells of cerebral cortex,
300
descending
345, 362
3"
ganglionic arteries. 19, 21 Postero-medial column (Golli), 300 Postero-median ganglionic arteries,
165
decussation, 291,
311
272,
15, 18, 19
Precentral lobule of island, 72 sulcus of cerebellum, 247, 248,
420
of cerebrum, 59, 410, 414
motor paths, 366 tract, 103, 148, 159, 226, 277, 294, 355, 358 Pyramis cerebelli, 87, 254
medullae oblongatae, 294 vermis, 254
INDEX.
Quadrangular lobule 243, 250 251: 255, 420 Quadrigeminal bodies, 4, 146, 147,
163
colliculus,
i.,
449
occipito-thalamica
104,
(optic),
Rhinencephalon, 28, 77, 97, 98, 406 Rhombencephalon, 31, 33, 38, 242, 266, 284 Rhomboid lip, 285, 422, 423 Rolandic angle, 56 Rolando, central sulcus of, 56, 57, 61, 410 gelatinous substance of, 339, 341
of fourth ventricle,
3M
Roof-plate, 400, 416, 422, 422, 427 Roots of optic tract, 83, 147, 153 of spinal nerves, 349, 362 of twelve cerebral nerves, 42 Rostrum of corpus callosum, 95, 108,
1 20
Radix anterior
363
of spinal nerve,
349,
349,
tract,
3'
305,
3"i
34i, 358,
37. 3?i
Salivary nucleus, 283 Schultze, comma tract
183
visual center, 81, 98, 181 Recessus triangularis, 128, 136,
341, 361
vestibular
Schwalbe's
dorso-medial
137
Red
nucleus, 101,157,159,204,217 Reflex arcs, 390, 391, 392 connections of abducent nucleus, 282 of cochlear nucleus, 310, 323 of facial nucleus, 282 of genetic nuclei, 325 of motor nuclei in medulla, 305 of nucleus alae cinerea?, 306
of olfactory nucleus, 324 of trigeminal nuclei, 281, 307 of vestibular nuclei, 308, 323,
nucleus, 280, 307 Schwann's sheath, 168 Second cerebral vesicle, 35 cervical nerve, 3
Secondary brain
sulci
vesicles, 402,
403
and
fissure, 410,
411, 414
mechanism
paths, 390
389.
of cord, 343
Semilunar ganglion (Gasseri), 45 nucleus, 278 Sense of touch, 377 Sensory aphasia, 241 cerebral nerves, 320 conduction paths, 377 paths, 229, 377
projection fibers, 231 roots of cerebral nerves, 320 of spinal nerve, 349, 363, 364 root of trigeminal nerve, 307
tract Ciaglinski,
326,
327,
329
of dura mater, i, 326, 329 of pia mater, 9, 327, 329 Respirator}' reflex, 393
350
Septo-marginal tract, 345, 361 Septum pellucidum, 30, 87, 95, 97,
288,
295, 297,
in, 115, 121, 123, 209, 235, 4i3. 4iS Seventh and eighth nerves, 3
45
INDEX.
Stereognostic center, 64, 98, 182,
Stilling's nuclei,
183
348
237
of),
75
Strand cells, 340 Stratum cinereum, 256 gangliosum, 256, 257 granulosum, 256, 258 griseum centrale, 152, 157, 159
221, 222 of superior colliculus, 163, 219 intermedium, 208, 225 interolivare lemnisci, 292
superior, 3 rectus, 2
sagittalis inferior, 2 superior, 2
transversi, 2
219
of thalamus, 140, 208 Stria intermedia, 77, 78, 267
lateralis, 78,
Sixth nerve, 3
ventricle,
335, 337
267
medialis, 77, 78, 267 medullaris thalami, 87, 91, 95, 135- *37> 212, 239, 239
terminalis,
124,
Striae
Somrcsthetic area, 64, 98, 181 Special nuclei of medulla, 302, 310
sensations, 383 sense fibers of internal capsule, 104, 115, 226 nuclei, 322
101, 109, 115, 119, 125, 207, 239, 412 medullares, 149, 270, 290, 309
olfactoriae, 78,
267
Striato-thalamic fibers, 207 Structure of arachnoid, 8, of cerebrum, 164 and relations of dura mater, and relations of pia mater, 9
Subarachnoid
rivulets,
426
ganglion. 334, 364, 401, 428 neurones, 165, 400, 401
reflexes,
390
trigeminal nerve,
278,
tract
of 301,
423, 424
91, 95
256
305,
3H,
Spino-vestibular tract, 357 Splenium of corpus callosum, 75, 95, 105, 108, 113, 109, 121,
173, 339
131
Spongioblasts, 398, 399 Stalk of pineal body, 87, 149 Stellate cells of cerebellum, 257, Stem of fissura cerebri lateralis,
Stereognosis, 377
259 57
101, 145, 146, 151, 157, 159, 161, 221 perforata anterior, 38, 75, 79, 200 posterior, 38, 75, 151 reticularis, 291, 295, 297, 302, 303- 305, 423 spongiosa, 339
INDEX.
Sulci, 52
451
Sulcus temporalis inferior, 70, 80, 86 medius, 61, 63, 70 superior, 61, 63, 70
vallecukc, 251, 252 Superciliary border, 39, 52 Superficial annectant gyrus, 64 layer of cerebellar cortex, 256
420
of convex surface of cerebrum, 52 of lower surface of cerebellum,
252, 253 of lower surface of cerebrum, 73 of medial surface of cerebrum,
85
275
Superior cerebellar artery, 15, 27 peduncle (see brachium conjunctivum) cerebral veins, 22
colliculus
upper surface of cerebellum, 248, 249 Sulcus basilaris, 153, 272 centralis (Rolandi), 39, 53, 56, 57, 61, 41 insulae, 65, 72 cinguli, 25, 86, 91, 95, 410 circularis insuhe, 70, 71, 75, 125
of
quadri-
91
39,
59,
60,
61,
209
sulcus,
hypothalamicus, 91, 139, 405, 4iS intermedius posterior, 335, 338 interparietalis, 61, 63
lateralis anterior, 41, 286,
39, 57,
59,
414
glossopharyngeal ganglion, 46
338
lamina of internal capsule, 100, 113, 115, 226 longitudinal fasciculus, 240 medullary velum, 13, 87, 146, 147, 149, 245, 246, 247,
418
252,
sig,
262, 263,
275
308
61, 86
uccipital gyrus,
75
olivary nucleus,
280
parietal lobule, 43, 61, 63, peduncle of cerebellum, 245
61
quadrigeminal
colliculus,
137,
61, 63
204,
superior, 61, 63
420
surface of cerebellum, 243, 248
of temporal lobe, 70 temporal gyrus, 61, 70, 71 sulcus. 57, 63, 70, 410 Supero-medial border, 39, 52
post decli vis cerebelli, 249 postnodularis cerebelli, 252 postpyramidalis cerebelli, 253
rhinalis,
80
subparietalis, 86,
91
452
INDEX.
Thalamo-striate
fibers,
Supracallosal gyrus, 97, 107, 414 Supramarginal gyrus, 43, 61, 63, 67 Supraradiary zone of cerebral cortex,
208
Third cerebral
vesicle,
35
37, 101, in,
nerve, 3, 4, 5, 42
ventricle, 9, 30, 34,
184 Surface of cerebellum, 248 of cerebrum, 34, 51 Surfaces of medulla oblongata, 286 of mid-brain, 144 of pons, 266
of spinal cord, 337 of thalamus, 140
127, 137, 145, 209 Threshold of island, 72. 77 Tigroid bodies, 167 Tonsil of cerebellum, 245, 252, 254,
420 Torcular herophili, 2 Tracing of impulses, 366 Tracts of fibers in tegmentum, 156
of spinal cord,
429
Taenia semicircularis, 115, 119, 125
(see stria terminalis) terminalis, 119 ventriculi quarti, 149, 270, 297
Tractus cerebro-cortico-pontalis frontalis, 103, 151, 225 temporalis, 100, 148, 225 cerebro-spinalis pyramiclalis, 226
intermedius, 100, 148, 225 olfactorius, 77, 78
optici, 83 solitarius,
Tapetum,
123, 124,
Tegmentum,
of mid-brain, 30, 75, 109, 132, 146, 152 Tela chorioidea ventriculi quarti, 10
a,
tertii,
422, 423
spinalis n. trigemini, 277, 278, 288, 295, 297, 301, 305,
10
3ii
Transverse fibers of medulla, 291 of pons, 271 of spinal cord, 351 fissure of cerebellum, 36 of cerebrum, 35, 43, 53, 55, 109, 411 occipital sulcus, 43, 57> 68, 69 sinuses, 2, 3, 4, 5 temporal gyri, 70, 105, 121,
Temporal
91, 98
pole, $3, 70 Tempo ro -occipital gyrus, 81, sulcus, 81
97
125
70 Trapezoid body, 271, 272, 277, 280 Triangle of habenula, 140 of lateral fillet, 152, 164 Triangular tract of Helwig, 298, 305, 311, 312, 341, 345, 357 Trigeminal nerve, 41, 45, I53> 209,
sulci,
226
100,
Trigonum
(see
ala
Tertiary
sulci,
411
Thalamus, i., 28,30, 37,51, 101, 105, 119, 125, 139, 145, 147, 157, 208, 209, 416, 417 Thalamo-mammillary fasciculus (of Vicq d'Azyr), 145, 222, 237
124 habenuke, 140, 137, 243 n. hypoglossi, 149, 270 olfactorium, 47, 75, 77, 79, 200 Trochlear nerve, 45, 146, I47> 3 21 nucleus, 223
collaterale, 123,
INDEX.
Trolard's great anastomotic vein, 23 Truncus of corpus callosum, 95,
453
156,
108
341,
345, 343
Ventro-medial
cells of cord,
34i,345>
34
Ventricle of corpus callosum, 89 Ventricular gray matter of cerebrum,
acusticum, 149, 309 cinereum, 13, 289, 301 Twelfth nerve, 3, 49, 322 Types of neurones, 168
Uncinate fasciculus, 237, 239, 240 Uncrossed ponto-spinal tract, 280, 373
127
Vermis
Uncus hippocampi,
235
419
Unipolar neurones, 400, 401 Uvula vermis cerebelli, 87, 245, 247, 252, 254
superior, 244
Vagus nerve,
321
5,
46,
Vertebral artery, 3, 5, 14, 15, 27 ganglia, 401 Vessels of arachnoid, 9 Vestibular nerve, 280, 293, 320, 386
Vallecula cerebelli, 36, 244, 251 Valve of Vieussens, 246 Veins of cerebellum, 29 of cerebrum, 21
of medulla, 24 of pia mater, 13 of spinal cord,
37
path, 386 Vestibulo-spinal reflex, 393
Vicq
Vicq d'Azyr (foramen of), 153 d'Azyr's fasciculus thalamomammillaris, 85, no, 222,
331, 332
237
Vieussens's medullary velum, 146
valve, 246
Velum
Vinculum,
a, cerebelli,
250
Vision, center for macular, 67 Visual center (receptive), 81, 98, 192 memory center, 67, 69
magna, 21
media, 23
terminalis, 235 Venae cerebelli inferiores, 29
superiores, 29
cerebri, 21
Wallerian degeneration, 171 White anterior commissure, 335, 351 matter of cerebellum, 262 of cerebrum, 224 of medulla, 290
of pons, 270 of spinal cord, 351 of thalamus, 208 Worm of cerebellum, 243 Writing center of Gordinier, 175, 178,
externae, 22
inferiores,
23 mediales, 22 superiores, 22
spinales extreace, 332
internae, 332 Ventral cochlear nucleus, 309 longitudinal fibers of pons, 272 stalk of thalamus, 100, 212, 226,
182
Zonal layer of cerebral cortex, 184 Zones of embryo, ventral and dorsal, 399
232